EDITOR-IN-CHIEF

Dr. José H. Leal

The Bailey-Matthews Shell Museum

3075 Sanibel-Captiva Road

Sanibel, FL 33957

MANAGING EDITOR

Linda Kramer

The Bailey-Matthews Shell Museum

3075 Sanibel-Captiva Road

Sanibel, FL 33957

EDITOR EMERITUS

Dr. M. G. Harasewych

Department of Invertebrate Zoology

National Museum of

Natural History

Smithsonian Institution

Washington, DC 20560

CONSULTING EDITORS

Dr. Riidiger Bieler

Department of Invertebrates

Field Museum of

Natural History

Chicago, IL 60605

Dr. Arthur E. Bogan

North Carolina State Museum of

Natural Sciences

Raleigh, NC 27626

Dr. Philippe Bouchet

Laboratoire de Biologie des

Invertébrés Marins et Malacologie

Muséum National d'Histoire Naturelle

55, rue Buffon

Paris, 75005 France

Dr. Robert H. Cowie

Center for Conservation Research

and Training

University of Hawaii

3050 Maile Way, Gilmore 409

Honolulu, HI 96822

Dr. Robert T. Dillon, Jr.

Department of Biology

College of Charleston

Charleston, SC 29424

Dr. Eileen H. Jokinen

8234 E. North Shore Road

Sault Ste. Marie, MJ 49783

Dr. Douglas S. Jones

Florida Museum of Natural History

University of Florida

Gainesville, FL 32611-2035

Dr. Harry G. Lee

1801 Barrs Street, Suite 500

Jacksonville, FL 32204

Dr. Charles Lydeard

Biodiversity and Systematics

Department of Biological Sciences

University of Alabama

Tuscaloosa, AL 35487

Bruce A. Marshall

Museum of New Zealand

Te Papa Tongarewa

P.O. Box 467

Wellington, NEW ZEALAND

Dr. James H. McLean

Department of Malacology

Natural History Museum

of Los Angeles County

900 Exposition Boulevard

Los Angeles, CA 90007

Dr. Paula M. Mikkelsen

Department of Living Invertebrates

The American Museum of Natural

History

New York, NY 10024

Dr. Diarmaid O Foighil

Museum of Zoology and Department

of Biology

University of Michigan

Ann Arbor, MI 48109-1079

Dr. Gustav Paulay

Florida Museum of Natural History

University of Florida

Gainesville, FL 32611-2035

Mr. Richard E. Petit

P:Ov Box 30

North Myrtle Beach, SC 29582

Dr. Gary Rosenberg

Department of Mollusks

The Academy of Natural Sciences

1900 Benjamin Franklin Parkway

Philadelphia, PA 19103

Dr. Angel Valdés

Department of Malacology

Natural History Museum

of Los Angeles County

900 Exposition Boulevard

Los Angeles, CA 90007

Dr. Geerat J. Vermeij

Department of Geology

University of California at Davis

Davis, CA 95616

Dr. G. Thomas Watters

Aquatic Ecology Laboratory

1314 Kinnear Road

Columbus, OH 43212-1194

Dr. John B. Wise

Department of Biology

College of Charleston

Charleston, SC 29424

SUBSCRIPTION INFORMATION

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for institutions. Postage outside the

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air mail. All orders should be

accompanied by payment and sent to:

THE NAUTILUS, P.O. Box 1580,

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22.331

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THE NAUTILUS (ISSN 0028-1344)

is published quarterly by The Bailey-

Matthews Shell Museum, 3075

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33975.

Periodicals postage paid at Sanibel,

FL, and additional mailing offices.

POSTMASTER: Send address

changes to: THE NAUTILUS

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PRE JNA th

Volume 120, Number 1

May 50, 2006

ISSN 0028-1544

CONTENTS

José H. Leal Celebrating a long life: The Nautilus turns 120! 00. 00 ee |

Nikolaus Malchus Amended description of the arcoid bivalve Philobrya brattstromi

Soot-Ryen, 1957, from Chile... 0.0.5 620. eek seed eee aed ee 8

Claude Vilvens Descriptions of Otukaia crustulum new species (Gastropoda: Trochoidea:

Javier Sellanes Calliostomatidae) and Margarites huloti new species (Gastropoda:

Trochoidea: Trochidae) from a methane seep area off Chile... 0.0.0.0... 15

Fred G. Thompson A new land snail of the genus Humboldtiana (Gastropoda: Pulmonata:

Humboldtianidae) from Nuevo Le6n, Mexico... 0.000.006 02.0002005 21

Fred G. Thompson Two new land snails of the genus Humboldtiana (Gastropoda: Pulmonata:

Omar Mejia Humboldtianidae) from Chihuahua, Mexico... ee 25

W. Wyatt Oswald Mollusks in a Holocene lake-sediment core from the Arctic Foothills of

northern Alaska 2... ee ns 30

Michelle L. Boudreaux Mytella charruana (Bivalvia: Mytilidae): a new, invasive bivalve in

Linda J. Walters Mosquito Lagoon, Florida ......... 0.0.0.0. ccc eee eee eee 34

NOYGCE: 5 ic eee eee 15 6H REG BG O64 Oe Sb eRe CE OS Oe Eas EA ben Sh awe KE eee P eae gas 37

MBLWHCI Library

JUN 1 2 2008

WOODS HOLE

Massachusetts 02543

THE NAUTILUS 120(1):1-7, 2006

Page ]

Celebrating a long life: The Nautilus turns 120!

José H. Leal

The Bailey-Matthews Shell Museum

P.O Box 1580

Sanibel FL 33957 USA

[email protected]

The Nautilus is tuming 120 years this year and I feel

extremely honored and lucky to be wr iting this note to

celebrate the occasion! The Nautilus is the one of the two

oldest English-language malacological journals, second

only to the Journal of Conchology in longevity in this

category. My goal is to briefly showcase, particular ly to

readers from younger generations, some highlights of the

journal’s ev olution ; and of the work of its Scions:

AN INAUSPICIOUS BEGINNING?

The Nautilus started in Philadelphia in July, 1886, as The

Conchologists’ Exchange (TCE). William D. Averell

(1853-1928), a shell ee from Chestnut Hill, Philadel-

phia, created TCE as a small publication aimed at “giving

information of vital interest to the student of Mollusea®

(Averell, 1S86a).

In November of that same year Averell (1886b) ob-

served: “It is our desire that each number shall be an

improvement upon its predecessor; . It is inevitable

that new discoveries will continue to be made in our

chosen field of research from now until the end of time,

as vast areas have yet to be fully explored and their trea-

sures described; and, again numerous species are rapidly

becoming extinct, while others are scarcely known or

have never had the light of discovery cast upon them.”

Averell’s words prov al. to be mostly prophetic in the long

run. But the immediate future of TCE was uncertain.

Starting in May, 1858, after 21 months of publication, the

periodical went out of circulation for one year,

Former editor R. Tucker Abbott in his dedication of

the 100 volume (1986) of the journal observed that

“The Nautilus had a very inauspicious beginning. . . . Vol-

ume 1, number 1, was merely a postcard sent out i 500

people announcing that the annual subscription price . . .

would be all of 25 cents.” The first two volumes each

consisted of a few pages published monthly in a duo-

decimo (12 mo) size (16.5x14 cm). The contents of TCE

covered specimen exchange notices for collectors, short

articles about collecting and preservation of mollusks,

news, and brief narratives of field trips, sometimes to

distant and exotic places such as Tampa, Florida.

PILSBRY TO THE RESCUE

Young conchologist and TCE contributor Henry Augus-

tus Pilsbry (1862-1957) (Figure 1) was hired as conser-

vator of the C onchological Section of the Academy of

Natural Sciences of Philadelphia (ANSP) in 1888, a few

weeks after he turned 25. After a year’s lapse and fol-

lowing negotiations between Pilsbry and Averell, in May,

1889, "TCE became The Nautilus (volume 3, number 1),

with Pilsbry as its editor. The journal size was adjusted to

crown octavo (about 19x13 em).

In his first number Pilsbry changed the physical for-

mat of the journal from double- column to single-column.

In addition, he prominently displayed, starting on page 2,

a brief technical article by W. H. Dall (1889) on the

anatomy of the gastropod Trochus infundibulum. Dall’s

paper, which differed considerably in style and content

from the average note and exchange notice previously

published in TCE, marked the beginning of Pilsbry’s

shift in orientation of the journal. That change gave The

Nautilus a new lease on life.

In all likelihood, Pilsbry assumed from the start that

the reorganization of TCE as the more science-oriented

The Nautilus was the natural thing to do. In the very first

sentence of the Introduction to volunie 3 (Pilsbry, 1589)

the 26 year-old editor observed: “The publishers of The

Nautilus feel that no explanation of their [objective | in

offering this journal to the scientific public is necessary.

On the same page, he indicated that it was his w ish to

have m: alacological papers concentrated in a few special-

ized pe fpdicals rather than in “the pages of innumerable

[general science] journals.” By “establishing journals de-

voted to special branches of science” such as malacology,

it would be possible to “limit by some means the number

of publications in which a certain subject is likely to be

treated upon.

Possibly anticipating a decline in interest from sub-

scribers, Pilsbry proposed that, upon the transition from

TCE to The Nautilus, “All subscribers . . . will be allowed

one insertion of twenty-five words in the Exchange Col-

umn, free of charge.” The appeal to a diverse audience

was epitomized in a brick-colored, promotional 15.28.2

cm flyer (Figure 2) published sometime in the early

1890s. The flyer prottered: “Its scope is broad, including

Page

THE NAUTILUS, Vol. 120, No. 1

Henry Augustus Pilsbry, 1862-1957; editor 1S589—

1957. Photo The Noutilss archives.

Figure I.

articles on recent and fossil shells... Notes... showing

the drift of opinion on conchological subjec ‘ts, are a valu-

able feature”. In addition, the footer of the flyer an-

nounced: “Exchange column free to subcribers.” The

subscription price was $1.00 per year, $1.12 to foreign

countries.

W. D. Averell remained as business manager of The

Nautilus after the transition until the end of 1890. Ac-

cording to Baker (195S8b), “His [Pilsbry’s] one year’s

association with allan D. Averell as its business man-

ager was none too happy and, at the end of one volume,

Averell was replaced by ... Charles W. Johnson . . .” In

a brief obituary of Avere ‘IL Pilebivt (1928) wemaeds “He

was deeply interested in conchology, but made no per-

manent collection. He was chiefly ape as a dealer in

shells and in connection with his little journal, which

proved to be highly useful for bringing mete »r the con-

chologists and eolle xctors of the late ’80s.”

Pilsbry’s tenure as editor of The Nautilus equaled in

length and paralleled his long and productive career as a

scientist (see Baker, 1958a); he edited the new version of

the “little journal” between 1888 and 1957, a 69-year

editorial stint probably unmatched in duration in the

world of malacological publications. Pilsbry unquestion-

ably consolidated the reputation of The Nautilus as a

THE

NAUTILUS

A MONTHLY

DEVOTED TO THE INTERESTS OF

CONCHOLOGISTS.

EDITORS AND PUBLISHERS:

s Conservator Conchological Section

H. a. PILSBRY, { Academy Natural Sciences, Philadelphia.

Ya Curator of the Wagner Free Institute of Science,

C2W.< JOHNSON { Philadelphia,

$1.00 per Year. $1.12 to Foreign Countries.

10 ets. a Copy.

Tue Navtitus is the only magazine in

America devoted to the study of Mollusks.

Its scope is broad, including articles on recent

and. fossil shells, often illustrated; on the

anatomy and classification of szollusca; and

papers of popular interest dealing with experi-

ences of collectors in the field. Notes on

current publications at home and abroad,

showing the drift of opinion on conchological

subjects, are a valuable feature. /udispensadle

to the Conchologist and Paleontologist.

Exchange column free to subscribers.

Figure 2. Promotional flyer, circa early ]S90s. Note emphasis

on scientific aspects combined with appeals to the non-

professional audience of the journal. The line drawing portrays

North American land snail Xolotrema denotatum (Férussac,

1821). The same drawing appeared earlier, as Helix palliata

Say, in Binney (1857), Binney and Bland (1869), and Teator

(1890).

cutting-edge publication dealing with the many facets of

20" Century malacology. Pilsbry’s biographers com-

mented about his fortainee and love for The Nautilus to

the very end. Daughter Elizabeth Pilsbry (1958) ob-

served: “Of Father, at nearly 95, one truly may say: ‘He

died young.”

EARLY BUSINESS MANAGEMENT

Pilsbry was aided for most of his career as editor of The

Nautilus by Charles Willison Johnson (1863-1932) (Fig-

ure 3), Johnson was curator at the Wagner Free Institute

J. H. Leal, 2006

Page 3

2: business

Photo Museum Comparative Zoology,

Figure 3.

manager 1590-1932.

Harvard University.

Charles Willison Johnson, 1863-193

of Philadelphia between 1588S and 1903; from 1903 to his

death in 1932 he was principal curator of the Boston

Society of Natural History. Early in his career as curator

at the Wagner Institute, Johnson replaced W. D. Averell

in 1S90 as business manager of The Nautilus.

New England malacol ygist Arthur Fairfield Gray

(Gray, 1932) remarked about Johnson’s role with The

Nautilus: * (Pik yry's] choice was most fortunate; Johnson

established the most cordial relations with subscribers

and contributors. During extended periods, when Dr.

Pilsbry was away on collecting trips, he acted as editor as

well as business manager.” Gray commented on the per-

sonal relation between the two editors: “In those cays

The Nautilus was a monthly. This co-partnership and

intimate friendship continued for forty-two years, until

terminated by Mr. Johnson’s death.” Figure 4 portrays a

sample of Johnson's ledger work in the early 20" Cen-

tury as business manager of The Nautilus.

Horace Burrington Baker (1889-1971) (Figure 5),

Professor of Zoology at the University of Pennsylvania

and a specialist in terrestrial g gastropods, became business

manager of the journal upon the death of Charles W.

Johnson in 1932. Baker served as an associate editor for

25 years. Abbott and Wurtz (1971) remarked that Baker

“was one of America’s outstanding land mollusk anato-

mists.” He was also an excellent illustrator, as attested by

the exquisite anatomical drawings in his works.

LIFE AFTER PILSBRY

After Pilsbry’s death in December, 1957, Baker pub-

lished a memorial issue (volume 71, number 3) appro-

priately named “The Pilsbry Nautilus”. In that issue,

Baker (1958b) observed: “At least twice during Septem-

ber ee October, 1957, Dr. Pilsbry implored me to make

sure that The Nautilus would continue, and solemnly was

promised that it would.”

And continue it did, Beginning in April, 1958, with

volume 71, number 4, The Nautilus was edited by a tri-

umvirate of malacologists led by Baker, now editor-in-

chief of the journal. Baker outlined the opel ee plan

for the journal following Pilsbry’s death: Charles B.

Wurtz (1916-1982) (Figure 6) from Philade elphia, and

Robert Tucker Abbott (1919-1995) (Figure 7), newly

hired to fill the Pilsbry C ie of Malacology at the ANSP,

“will join the staff as junior editors with the understand-

ing that, if at any future time either should cease to be a

resident of the Philadelphia area, this would be consid-

ered as a tendered resignation” (Baker, 195Sb).

R. Tucker Abbott was initially in charge of marine

mollusks, Wurtz was responsible for mailing issues and

became editor for fresh water mollusks and e ecology, and

Bernadine B. Baker (“Mrs. Horace B. B.” \eeindadl to the

finances and subse os Baker added: “Since the Bak-

ers and The Nautilus can afford but one bank account,

checks may be made out as usual.” Between April, 1958,

(volume number 4) and April, 1972, (volume 85,

number 4), The Nautilus was subtitled “The Pilsbry

Quarterly aa to the Interests of Conchologists.” Be-

tween July, 1936 (volume 50, number 1) and April, 1972

(Volume 85, number 4), text on the second cover indi-

cated that The Nautilus was “... the official |... “an of-

ficial...” in the latter part of that period] organ of the

American Mal acological Union (now American Malaco-

logical Society).”

"Baker was editor-in-chief until 1968, and served as

Editor Emeritus until the time of his death in 1971.

Abbott and Wurtz (1971) reminisced: “For 40 years he

gave unstintingly of his time as business manager and

editor of The Nautilus.”

TUCKER'S TENURE

R. Tucker Abbott had been acting editor of The Nautilus

since H. B. Baker had his first flucey of health compli-

cations in 1968. He left ANSP in that year to occupy the

newly created Du Pont Chair of Mal: icology and serve as

assistant director of the Delaware Museum of Natural

History. Upon Baker's death in March, 1971, he formally

became editor-in-chief of the journal, helped by Charles

Wurtz and with Mrs. Baker as business manager.

Tucker edited The Nautilus from 1968 through L985,

making several changes in the format and editorial policy

that basically shaped the journal as we know it ae

Most of these improvements took place in July, 1972

with the publication of volume $6, number 1. This was a

very hectic time in Tucker's life; among his many activi-

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Figure 4. First four pages of the ledger for 1915 in Charles W. Johnson’s longhand. Pages correspond to the initial batch of

subscriptions for Volume 29. List of subscribers is a who’s who of American and international malacology of the period. Included are,

among others, John B. Henderson, Paul Bartsch, William F. Clapp, Calvin Goodrich, Edward S. Morse, M. M. Schepman, Don

Carlos de la Torre, William H. Dall, Frank C. Baker, and Victor Sterki. Some institutions listed (e.g., Museum of Comparative

ae Carnegie Museum, and University of Chicago) have been loyal subscribers through the entire 20'" Century and to the

present day.

ties then were the preparation of the trend-setting and Tucker made many changes in the appearance of the

classic second edition of his American Seashells and the journal. The size changed from crown octavo to

grand opening of the Hall of Shells at the Delaware 26.5x20.5 cm. He added a gray protective cover (each

Museum in May, 1972 (Harasewych, 1997). number in the preceding 85 years started with a contents

]. H. Leal, 2006

Page 5

Figure 5. Horace Burrington Baker,

1958— 1968. Photo The Nautilus archives.

Figure 6. Charles B. Wurtz (1916-1982) at the AMU meet-

ing at Haverford College in 1959. Photo from the American

Malacological Society archives.

1889-1971; editor

Figure 7. Robert Tucker Abbott, 1919-1995; editor 1968—

1985. Photo The Nautilus archives.

page of the same stock as the regular pages). An outline

of a slightly stylized nautilus shell in cross-section be-

came the logo (Figure 8), and the journal subtitle be-

came “A quarterly devoted to malacology and the inter-

ests of conchologists”.

Another advances introduced in the same number was

the addition of an editorial committee composed of 13

professionals representing a wide swath of American ma-

lacology. These were Arthur H. Clarke, Jr., William J.

Clench, William K. Emerson, Morris k. Jacobson, Au-

réle La Rocque, James H. McLean, Arthur S. Merrill,

Donald R. Moore, Joseph Rosewater, G. Alan Solem,

David H. Stansbery, Ruth D. Turner, and Gilbert L

Voss.

Last but not least, he shifted the publication of the first

number of each volume from July to January beginning

with volume 87 (1973): volumes now match calendar years.

Tucker's tenure as editor-in-chief spanned 17 years,

not counting his years as co-editor under H. B. Baker.

Tucker left the Delaware Museum in 1978 and, in his

own words, “moved The Nautilus, now under the juris-

diction of my American Malacologists, Inc., to Mel-

bourne, Florida, where it has been published for the last

six years [1979-1985, actually seven years]” (Abbott,

1986). In that period, Tucker's Wife Cecelia was the j jour-

nal’s business manager.

THE HARASEWYCH YEARS

In 1984 Tucker invited Miroslav George (Jerry) Ha-

rasewych (bor in 1949) (Figure 9), a former student

Page 6

THE NAUTILUS, Vol. 120, No. 1

Figure 8. Cover with new design introduced by Robert

Tucker Abbott in July, 1972.

and, at the time, a Research Fellow at the National In-

stitute of Mental Health, to join the editorial staff of The

Nautilus and to assist in various aspects of the production

of the journal. Following the publication of volume 100

of The Nautilus, Abbott retired as editor, to be suc-

ceeded by Harasewych, then a newly appointed Curator

of Marine Mollusca at the Smithsonian’s National Mu-

seum of Natural History. During its first hundred vol-

umes, The Nautilus was “owned” ‘by the editor. To com-

ply with the Smithsonian's requirement * ‘to avoid the

a for an appearance of a conflict of interest,”

ownership of The Nautilus passed from American Mala-

cologists to Trophon Corporation, a non-profit corpora-

tion established for the purpose during Harasewych’s

tenure as Editor.

Beginning with volume 101 (January 1987), Harase-

wych hecame editor-in-chief while Abbott continued as

associate editor until his death in 1995. Harasewych

again altered the format and contributed a new cover and

logo, designed and illustrated by Hal Lewis Design, Inc.,

of Philadelphia: this is the design still in use fodlay, in-

cluding Lewis's beautiful line drawing of a live Nautilus

pompilius on the cover, a nice improvement over the

previous layout.

In addition, Jerry further added to the professional

spin imparted by Tucker. He remarked (H: wrasewych,

1987) in the opening of volume LOL: “The Nautilus ‘will

Figure 9. M.G. (Jerry) Harasewych, bor 1949; editor 1985—

1998. Photo courtesy M.G. Harasewych.

continue to meet ever higher standards in the publication

of papers on all aspects of the biology and systematics of

mollusks.” At that time I was second: -year graduate stu-

dent at the Rosenstiel School in Miami and wondered

how hard it would be from then on to have a manuscript

accepted by the journal.

THE PRESENT

Jerry was editor until the baton was passed to me in May,

1998 (volume 111, number 3), after a brief stint as man-

aging editor from February, 1997 (volume 110, number

2) to February, 1998 (v olume 111, number 2). Tina

Yorgey was managing editor between March, 2001 (vol-

ume 115, aumber 1) and December, 2006 (volume 119,

number 4).

With the acquisition of The Nautilus by The Bailey-

Matthews Shell Museum in 1998, the journal became for

the first time owned by an academic organization. Tuck-

ers role as founding director of the museum until his

death in November, 1995, was ultimately instrumental in

the transter of The Nautilus to the organization.

The Nautilus results from a cooperative effort between

its staff, authors, reviewers, and subscribers. With your

help and support of future generations, I am certain that

the journal will thrive for at least another 120 years!

ACKNOWLEDGMENTS

Baseline biographical data for this note was gleaned from

the massive work by Coan, Kabat, and Petit (2006). For

key information I am grateful to Paul Callomon, M.G.

J. H. Leal, 2006

Page 7

Harasewych, Alan Kabat, Harry G. Lee, and Richard E.

Petit. M.G. Harasewych, Linda Kramer, Kimberly Nea-

lon, and Richard E. Petit critically reviewed the manu-

script.

LITERATURE CITED

Abbott, R. T. 1975 (ed.). The Best of The Nautilus. American

Malacologists, Greenville, 280 pp.

Abbott, R. T. 1986. Dedication of the 100" volume. The Nau-

tilus 100: 1-7.

Abbott, R. T. and C. B. Wurtz. L971. Horace Burrington Baker:

ISS9-LOTL. The Nautilus 85: 1-4.

[Averell, W. D.] 1SSGa. Salutatory, The Conchologists’ Ex-

change 1 (2): 1 (August 1SS6).

{Averell, W. D.| 1SS86b. Editorial. The Conchologists’ Ex-

change 1 (5): 1 (November 1SS6).

Baker, H. B. 195Sa. Henry Augustus Pilsbry: 1562-1957. The

Nautilus 71: 73-83, pls. 5-9.

Baker, H. B. 195Sb. The Pilsbry Nautilus. The Nautilus 71:

112-115.

Binney, A., 1857. The Terrestrial Air-Breathing Mollusks of the

United States and the Adjacent Territories of North

America. Volume 3. 40 pp. + 74 pls.

Binney, W. G., and T. Bland, 1869. Land and Fresh Water

Shells of North America. Part I. Pulmonata Geophila.

Smithsonian Institution, Washington, xii + 1-316 pp.

Coan, E. V., A. Kabat and R. E. Petit. 2006, 2.400 Years of

Malacology. 3"! edition. American Malacological Society,

664 pp. January 18, 2006. http:/Avww.malacological.org/

publications/2400_malacology.htnl

Dall, W. H. 1889. Notes on the soft parts of Trochus infun-

dibulum Watson with an account of a remarkable sexual

modification of the epipodium, hitherto undescribed in

Mollusca. The Nautilus 3: 2-4.

Gray, A. F. 1933. Charles Willison Johnson, 1863-1932. The

Nautilus 46: 129-134.

Harasewych, M. G. 1986. | Untitled introduction]. The Nautilus

101: 2.

Harasewych, M. G. 1997. The life and malacological contribu-

tions of R. Tucker Abbott (1919-1995). The Nautilus 110:

55-75.

Pilsbry, E. 1958. Collecting in Peru and Argentina. The Nau-

tilus 71: 116-iii [third cover].

Pilsbry, H. A. 1889. Introduction, The Nautilus 3: 1.

Pilsbry, H. A. 1928. William D. Averell. The Nautilus 42: 33.

Teaton [Teator], W. S. 1890. Collecting land snails in eastern

New York. The Nautilus 3: 109-110.

THE NAUTILUS 120(1):5-14, 2006

Page §

Amended description of the arcoid bivalve Philobrya brattstromi

Soot-Ryen, 1957, from Chile

Nikolaus Malchus

Universitat Autonoma de Barcelona

Dept. Geologia/Area Paleontologia

08193 Bellaterra, Catalunya, SPAIN

[email protected]

ABSTRACT

Cross-checking of the original material and labels for Philobrya

brattstromi nth the data from Soot-Ryen’s publications (1957,

1959) confirms that that author had de signated a holotype and

five paratypes by 1953. All specimens ouner than those from

station M24 and determined in the same year are thus excluded

from the type series. The correct height of the holotype is 5.5

mm and not 15 mm as originally indicated by Soot- Ryen (1957)

or 5.8 mm as he indicated ‘subseq uently (Soot-Ryen, 1959). The

prodissoconch of P. brattstromi is devoid of sculptural elements

(also at high magnifications) except for a raised, round-crested

shell margin as already described by Soot-Ryen. However, it

also has an incipient ‘dorso-central boss and well- developed

anterior and posterior wings. The mean length is 627 rm; the

L/H ratio of 1.5 is high when compared to eleven congeners for

which such measurements are available. Prominent morpho-

logical features of the postlarval shell are: (1) well-developed

posterior G1 teeth which considerably increase in length an-

tero-ventrally to ae Ae ligament and that disintegrate ven-

trally into pustule s; (2) four to five posterior tooth- like folds on

the inner margin Lee described by Soot-Ryen), which

correspond in number with ribs on the outer surface of valve;

(3) a non-flaky periostracum. The shell microstructure is linear

to complex crossed-lamellar with an inner prismatic layer. The

additional presence of a (vestigial?) outer prismatic layer dorsal

to the ligament area, the pustular disintegration of G1 teeth and

the presence of a dorso-central boss on the prodissoconch re-

semble these states in the Limopsidae. However, it remains

questionable whether these traits are inherited or convergent.

INTRODUCTION

Philobrya brattstromi Soot-Ryen, 1957, belongs to a

small but rather successful arcoid family, the Philobry-

idae, which radiated into about 85 extant species since

the Eocene; about 45 of these species are currently in-

cluded in the type genus Philobrya (Linse, 2004, pers.

comm.). All descriptive and phylogenetic work on Philo-

bryidae emphasizes the importance of prodissoconch and

postlarval hinge characters (e.g., Bernard, 1897; Dell,

1964; Hayami and Kase, 1993; Malchus and Warén,

2005; Tevesz, 1977). However, until now, details of these

characters and of the shell microstructure of Philobrya

brattstromi have not been examined using a high-

resolution microscope such as a scanning electron micro-

scope (SEM) and are therefore still poorly known (com-

pare with Soot-Ryen, 1957, 1959: pl. 1, fig. 6; Ramorino,

1968: pl. 5, figs. 2 and 4). In addition, Soot- Ryen’s origi-

nal publications cast doubts about the identity of ihe

holotype and thus also about the composition of ‘the type

series. The present contribution, therefore, aims to re-

describe the main shell features in a modem interpreta-

tional context and to clarify the type status of the speci-

mens.

MATERIALS AND METHODS

The material includes five lots from the invertebrate col-

lection of the Swedish Museum of Natural History in

Stockholm (SMNH). The type series from the collection

site “station M24” (SMNH 3894) consists of the holotype

with articulated valves (AV) and five paratypes (one dis-

articulated LV and RV, and three AV). Eight specimens

are from station M17 (SMNH 74831) (5 AV, 1 LV, 2 RV

fragments), two soft parts (no shell) from station M48

(SMNH 74832), two AV from station M92 (SMNH

74833), and one AV from station M103 (SMNH 74834)

Table 1). Specimens from stations M21 and M27 men-

tioned in Soot-Ryen (1959) are not in the collection.

Material from station M48 found in the collection is not

mentioned in the list of stations in Soot-Ryen (1959: 6).

One of the labels from SMNH 74834 indicates station

M104; this latter, however, is not mentioned in the list of

stations (1959). However, all other data on the label are

identical with those given for station M103 in that list

(Soot-Ryen 1959: 6) (Table 1). All specimens were pre-

served in ethanol, many of them with the periostracum

preserved to a large extent.

In addition to the literature, my own data on the fol-

lowing eleven species of Philobrya were used for com-

parisons: Philobrya crispa Linse, 2002; P. laevis Thiele,

1912; P. magellanica Stempell, 1899 ; P. cf. olstadi Soot-

Ryen, 1951, P. quadrata Ptetter, 1886 (in Martens and

Pfeffer, 1886): P. sublaevis P elke sneer, 1903: P. wande-

lensis Lamy, 1906; P. meleagrina Bernard, 1897 (type

N. Malchus, 2006

Page 9

Table 1. Data for the five lots of P. brattstromi from the Swedish Museum of Natural History (SMNH).

SMNH

number Station Date Locality Depth, substrate

3894 M24 16 Dee. 1948 Seno Reloneavi, S of Isla Guar, 41°44’ S, 70 m, sand with shells

72°45' W

74831 M17 14 Dec. 1948 Golfo de Ancud, Canal Calbuco, 41°46’ S 30 m, grey sand and small

73°06' W stones

74832 M4S 3 Feb. 1949 Seno Reloncavi, bay of Puerto Montt, 41°28" S, 30 m, ?

72°56’ W ,

74833 M92 3 May 1949 Golto de Ancud, Bahia Ilto, 41°53’ S$, 73°10' W 45 m, sand with dead algae

74834 M103 (M104?) 5 May 1949 Canal Chacao, N of Punta Soledad, 41°48’ S, 40 m, stones and polychaete

73°31' W

tubes

series) and P. olstadi Soot-Ryen, 1951 (type series); Phi-

lobrya ?new species from New Zealand identified as P.

meleagrina (SMNH specimens) (I also included com-

parisons with some species of Adacnarca and Lissarca).

This material is deposited in the British Antarctic Sur-

vey, Cambridge, UK, accessible through ongoing col-

laborative work with Katrin Linse on Southern Ocean

philobryids and the Swedish Museum of Natural pee

Stockholm, Sweden, examined during a research stay;

addition, I examined the type series of P. meleagrina

(Muséum national d'Histoire naturelle, Paris) and P. ol-

stadi (Zoological Museum, Oslo).

Specimens of Philobrya brattstromi were studied and

shell dimensions (length and height) measured using a

binocular microscope (x40 magnification). In addition,

the holotype was photogr aphed and measured under a

Leica MZ FLIII stereo microscope equipped with a Pla-

napo objective and a DC 500 digital camera. Both valves

of a disarticulated paratype ane. five specimens from lot

SMNH 74831 were examined and measured under

SEM. For this, specimens were cleaned in 70% ethanol

and some specimens also cleaned with commercial

bleach for dissolution of the periostracum. Specimens

Table 2. Dimensions of some prodissoconchs and adult shells

from the collection. (1) identifies adult shell dimensions with-

out periostracum. Abbreviations: AV, articulated valves; LV,

left valve; RV, right valve, n/d, not determined.

SMNH Prodissoconch Adult

number Notes L x H (pm) L x H (mm)

3894 AV (holotype) 659 « 414 49x55

LV + RV 649 x 409 DD

AV n/d 3.3 x 4.0

AV n/d 3.0 x 3.3

AV n/d 2.3 x 2.8

74831 RV fragment 620 x 442 n/d

AV (1) 591 x 403 26x 3.1

LV (1) 614 x 421 3.8 x 4.8

AV n/d 3.6 x 4.6

AV n/d 3.0 x 2.5

AV n/d 2.3 x 2.7

74833 AV n/d 4.0 x 5.2

AV n/d 3.3 x 3.8

74834 AV n/d 3.0 x 3.5

were mounted on aluminum stubs with adhesive carbon

pads and sputter-coated with gold. The microstructure

was described from broken shells of a disarticulated para-

type and one specimen from station M17 (SMNH 74831).

The lengths of the larval and postlarval shells were

measured as the longest anterior-posterior distance par-

allel to the straight hinge axis of the prodissoconch and

the height as the longest dorso-ventral distance perpen-

dicular to that axis (Figures 1, 2). Measurements of the

adult shell under light microscopy include the perios-

tracum; specimens measured under SEM lack the peri-

ostracum. Holotype dimensions were also determined

using the long axis of the shell (with periostracum) and

the longest extension perpe sndicular to the long axis (Fig-

ure 1). “This orientation appears to be equivalent to the

axes used by Soot-Ryen (1959). The umbonal angle is

defined as the angle enclosed by the tangents to the

postero-dorsal and antero-dorsal shell margins of the

postlarval shell (Figure 1).

HISTORICAL BACKGROUND

Tron Soot- Ry en was entrusted with the identification of

the bivalve fauna obtained during the Lund University

Chile Expedition 1948-1949. The original labels accom-

panying the Museum material eaeaié that Soot-Ryen

had established a holotype and paratypes and had deter-

mined all specimens now Seria in the collection by

1953. However, this is not evident from his original pub-

lication (Soot-Ryen, 1957), which provides only of a brief

preliminary diagnosis, making reference to the type lo-

cality and to a shell length of 15 mm, supposedly of the

holotype. A more comple te account of the taxon with a

taxonomic discussion and a list of collection sites we Is

published in 1959. The description (called “diagnosis”

therein) is identical to that from 1957 except that the

shell length of the now explicitly named holotype

rei to 5.8 mm (Soot-Ryen, 1959: 22-23, pl. 1, fig.

. \: This value matches the measurements obtained here

ae a slight deviation) whereas none of the collection

specimens is as large as 15 mm. Interestingly, Soot-Ryen

(1957: 3) cited a length of 5.8 mm for Lyonsia elegantula,

which directly follows the diagnosis of Philobrya

brattstromi in that paper. This v: alue changed to 15 mm

Page 10 THE NAUTILUS, Vol. 120, No. 1

umbonal angle

P-hinge axis Vv

Height

> Length al 250 um

external

folds

internal

folds

9g 10 11

Figures I-11. Definitions, terms, and general shell features. Scale bars = 1mm (where not indicated otherwise). 1. Outline of an

adult shell (left valve) indicating shell axes for measurements, 2. Outline of a left valve prodissoconch (from exterior) superimposed

on a dark circular background with specimen outline and circle sharing central point (see Figure 3). The size of such circle is defined

as the minimum size at which the shell contour becomes enclosed and the center of both shell and circle coincide. The acute angle

(< 90°) between dorsal and posterior shell margins (quadrants I and IT) is typical of many philobryid species (Malchus and Warén,

2005, fig. 5). 3. Prodissoconchs of right valve (top) and left valve of an articulated specimen (SMNH 74831). 4-6. Left valve,

prodissoconch, and right valve of holotype (SMNIL3894) (photos taken with optical microscope). 7-8. Left valve (left) and right valve

of two specimens (SMNH 74831). 9. Left valve of paratype (SMNH 3594), 10. Close-up of hinge of specimen in Figure 7 (SMNH

74531). LL. Detail of most ventral part of posterior GI teeth of Figure 9 showing disintegration of teeth into pustules (SMNH_ 3594).

N. Malchus, 2006

Page 1]

in Soot-Ryen (1959: 36) suggesting that the dimensions

of the two species were exchanged in the first publica-

tion. This inte 1 tation is supported by the fact that the

author described no other species with exac tly that size

in 1957.

SYSTEMATICS

Order Arcoida Stoliezka, 1S71

Family Philobryidae Bernard, 1S97

Description: — Small (<l0mm), brooding, with large

prodissoconch (length 40 1200. juin), postl: urval shell

more or less sig ore ly oval (antero-dorsal/postero-ventral

axis = long axis) _ Primary tooth series (generation | teeth,

G1) functional throughout adult life or reduced to pus-

tules in larger adults and then unlikely functional (e.g,

Philobrya sublaevis): primary, “larval” ligament (L1)

within primary pesiliter throughout adult life; with or

without a reduced number of generation 2 teeth (G2):

disjunct fibrous adult ligament (sensw Waller, 1990) lack-

ing and thus adult ligament not duplivincular, fibrous

ligament sublayer between LV and RV continuous, i.e.

without lamellar bridge (corroborating a communication

by Waller, 1989, to Carter, 1990: 195). (Description after

Bernard, 1897; Dell, 1964; Hain and Arnaud, 1992;

Keen, 1969: Tevesz, 1977; own observations; based also

on Malchus and Warén, 2005; Morton, 1978; Prezant,

1990):

Discussion: — Recent studies leave little doubt that the

postlarval first generation of hinge teeth and the liga-

ment/resilifer of philobryids are “homologous with fhe

larval to early postlarval hinge teeth and larval ligament/

resilifer of planktotrophic Pteriomorphia. Similarly, the

second generation of hinge teeth (e.g., Lissarca) is ho-

mologous with the adult tooth series as found in the

majority of arcoid bivalves (Malchus, 2004a, b; Malchus

and Warén, 2005: Prezant, 1990: and references

therein). The maintenance of functional Gl teeth and

primary ligament, the reduction of G2 teeth and the lack

ofa duplivincular adult ligament are deemed to indicate

paedomorphic evolution (Malchus and Warén, 2005;

Morton, 1978: Waller, 1990).

Genus Philobrya Carpenter, 1872

Description: — G2 teeth lacking, ligament and resilial

groove posterior ye longated, many or most species tear-

drop-shaped with a pointed umbo capped by prodisso-

conch, umbonal angle from 90° to 120° (between antero-

dorsal and posterodorsal shell margins; Figure 1), ante-

rior adductor muscle scar lacking. Periostracum not

always flaky. (After Tevesz, 1977; this study. )

Discussion: — The periostracum is not always flaky as

diagnosed by Keen (1969: N270). Philobrya differs from

Adacnarca by its more mytilid-like shape, anterior posi-

tion of its umbo, and a posteriorly elongated ligament

and groove. Both genera differ from Lissarca by their

lack of G2 teeth (Dell, 1990: figs. 41-45, 49, 54, 58).

Nicol (1966: 28) correctly described the hinge of Adac-

narca but incorrectly included Philobrya wandelensis in

that genus based on hinge similarities. More in line with

the results presented herein, Dell (1990: 26-27) assumed

that this similarity is superficial even though he inter-

preted the “vertical striae” as ligament pits rather than

G1 teeth. In fact, the similarity is not superficial but may

reflect a primitive trait (Dell, 1990: fig. 57 of a Dacry-

dium hinge; Malchus and Warén, 2005: figs. 1-2 of vari-

ous Limopsidae; among many others). For this reason,

Nicol’s concepts of Philobrya and Adacnarca are not ac-

cepted herein.

Philobrya brattstromi Soot-Ryen, 1957

Philobrya brattstr¢mi n. sp.: Soot-Ryen, 1957:

Philobrya brattstromi n. sp. (1957, p. 2): Soot-Ryen, 1959: 22,

pl. 1, fig. 6.

Philobrya brattstromi Soot-Ryen, 1957: Ramorino, 1968: p.

198, pl. 1, figs. 5-6, pl. 5, figs. 2 and 4,

Type Series: — By original designation, the type series

includes only the specimens from station M24 (SMNH

3894) even though the author determined all specimens

of the Sellection: in the same year (International Com-

mission on Zoological Nomenclature [1999]: Articles

2.4.1 and 72.4.6.).

Holotype: — The holotype and paratypes were desig-

nated by the author. The holotype is complete with a rather

well preserved prodissoconch, articulated and closed shell

with a protruding byssus and an almost entire, transpar-

ent, and non-flaky periostracum. The periostracum ex-

tends up to about 0.9 mm.beyond the shell and forms 11

larger radial ridges with hairy bristles (Figures 4-6).

This specimen ee ser with the figure given by

Soot-Ryen (1959, pl. 1, fig. 6) (except at the RV was

found broken antero- eontrally). However, the dimen-

sions found here only coincide with those of Soot-Ryen

(1959), if measurements (1) include the periostracum,

(2) his “length” refers to the longest dorsal-ventral axis

(height in the present terminology) and (3) his “height”

(length here) is measured pe rpendicular be that dorsal-

ventral axis. Thus measured, the shell is 4.1x5.5x2.1 mm

(length x height x width) versus 4.4%5.8x2.3 mm given by

Soot-Ryen. U sing the straight larval hinge as a reference

(as is done throughout he re), dimensions are 4.2x4.9 mm

(LxH) (Figure 1).

Year of Publication: — Although the preliminary diag-

nosis of P. brattstromi from 1957 is rather rudimentary,

that publication appears to fulfill the requirements of the

International Commission of Zoological Nomenclature

(1999; particularly Articles 11, 13) and 1957 should be

considered as the valid year of publication.

Original Description (Soot-Ryen, 1957: 2): “Shell

small, oblique ovate, white, w ith faint r: adiating ribs. Pe-

riostracum yellow with 9 to 13 ribs formed by protruding

hairs over the ribs of the shell, continuing beyond the

shell margin. Shell with numerous fine pittings inside the

pallial line. Anterior margin very short with rounded dor-

Page 12 THE NAUTILUS, Vol. 120, No. 1

250 um 250 um 250 um

12 13 14

250 um 250 um 250 um

15 16 17

25 um

——

21 22

Figures 12- | Prodissoconch, central hinge, and microstructure of postlarval shell. 12. Central hinge of left valve (SMNH 74831;

see Figures 7, 10). 13. Central hinge of left val e of paratype (SMNH 3894; see Figure 9). 14-15. Cental hinge of right valve of

paratype, ari sriorly broken. Note Gack of tooth that would fit in the depression of the opposite valve (Figure 13), Figure 15 shows

rie saris crested, raised margin of the prodissoconch (SMNH 3894). 16. Prodissoconch of left valve of paratype (same as Figures

13, SMNH 3894). 17. Right-valve prodissoconch, same specimen as Figure 3 (SMNH 74831). 18-19. Dorsal-ventral fracture

abs showing central portion of right valve paratype with pores. 19. Detail showing outer cross-lamellar, middle complex

cross-lamellar, and inner prismatic (same specimen as Figures 14-15) (SMNH 3894). 20-23. Dorsal-ventral fracture surfaces of a

specimen from SMNH 74831. 20. Middle of the shell with pores penetrating each other. 21. F cai near ventral shell margin,

showing low penetration angle (about 35°) of pores with respect to inner shell margin (to the right). 22. Fracture through ambonal

area. 23. Detail of Figure 22, showing complex cross-lamellar structure covered by very thin outer prismatic shell layer (white

arrows).

N. Malchus, 2006

Page 13

sal angle, linearly descending to the rounded ventral

margin which continues to a rounded posterodorsal

angle. Dorsal margin slightly convex or straight. Prodis-

soconch with raised margins and without visible sculp-

ture. Hinge with anterior tooth-like projection corre-

sponding to a depression in the other valve. From the

anterior part of the prodissoconch a rather broad, cross-

lined part extends backwards below the resilium, which

starts below the middle of the prodissoconch and con-

tinues backwards and inwards to the middle of the hinge

plate. Above and behind the ligament the cross-lining

continues to the end of the hinge plate, before which

there are 4 to 5 oblique, tooth-like folds in the margin.

Adductor scar placed considerably within the pallial Tae

in the posteroventral part. Mantle mar ein with brownish

pigment, but lacks eyes. Length 15 mm.” (Should read

5.8 mm, see above).

Amended Shell Description: — The prodissoconch is

moderately elevated with a round-crested rim and steep

ramp tow ards the postlarval shell, the surface is smooth,

except for posterodorsal and anterodorsal wings marked

by growth increments and a small darséesatcel bump-

like elev ation (boss); mean prodissoconch size is 627x415

uum (LxH) (5 specimens), with a L/H ratio of 1.5 (Fig-

ures 2—3, 12-13, 15-17)

Postlarval G1 teeth (“cross-lining” of Soot-Ryen) well

developed; ventral to the long ligament and groove, the

teeth of the posterior row grow considerably in length,

then the row tapers and teeth disintegrate into irregular

pustules ( (Figures 10-11). The “anterior tooth-like pro-

jection’ mentioned by Soot-Ryen does not correspond to

a depression in the other valve; instead, both valves show

a depression directly ventral to the anterior row of G1

teeth (Figures 12— 14). The posterodorsal radial folds on

the inner shell margin correspond to an equal number of

radial ribs on the outer surface, which are slightly more

prominent than the other radial ribs (Figures 7-9).

Most of the shell has a linear crossed-lamellar struc-

ture (CL), further internally the structure is also complex

(CCL); the innermost layer is prismatic (P) with up

to 10 pm thickness (Figures 15-19). An outer prismatic

layer 1 wm thick was only observed in a zone dorsal to

the ligament (Figures 22— 23). Numerous, mainly straight

shell pores ( (“fine pittings” of Soot-Ryen) penetrate the

inner (depositional) surface but not all extend to the

outer shell surface, some occur in the prodissoconch.

Pores are distributed almost homogenously over the

depositional surface within the pallial line. Diameters

range from 1.5 to 2.5 wm within the shell, the funnel-like

entrances have inner diameters of 2 to 4 wm. Penetration

angles are mostly perpendicular to the (inner) central

shell surface but about 35° near the shell margins; rami-

fications were not observed but pores occasionally pen-

etrate each other (Figures 18-21). Adult shell size is up

to about 5x6 mm (LH).

Distribution: Golfo de Ancud (41°44 S) to Bahia de

Valparaiso (33° S) along the Chilean coast (Soot-Ryen,

1959; Ramorino, 1968).

Discussion: — Philobrya brattstromi is an easily recog-

nized species given the combination of (1) a smooth pro-

dissoconch with incipient dorso-central boss, well devel-

oped anterior and posterior wings, and mean le cue of

627 ym and relatively high L/H ratio of about 1.5,

well developed posterior G1 teeth reaching far a

enhancing in length anteroventral to the ligament and

disintegrating into pustules, (3) the number of posterior

radial ridges which correspond in number with folds on

the inner shell margin which is otherwise smooth, and (4)

a non-flaky periostracum.

Most of the other shell characters described here are

either typical of the family Philobryidae or of the genus

Philobrya (see respective descriptions, above), and shell

pores are characteristic for probably all Arcoida (Taylor,

Kennedy and Hall, 1971; Malchus and Warén, 2005;

Waller, 1980). Similarly, an approximately homogenous

distribution of pores appears to be common for most

arcoids even though they occur concentrated in radial

rows in some (or all?) Adacnarca species.

Accordingly, it is significant that characters such as the

presence of an incipient dorso-central boss, the disinte-

gration of G1 teeth and the presence of remnants of an

outer prismatic layer are similar to those in oe

(Carter, 1990: 194: Malchus and Warén, 2005). G1 dis-

integration is also common in other species of Philobry

and although a small dorso-central boss appears to be

rare in this genus, this feature appears to develop into a

prominent cone in the philobryid genera Cratis and Cosa

(e.g. Hayami and Kase, 1993). The distribution of these

characters is consistent with the generally assumed an-

cestor-descendent relation between Limopsidae and Phi-

lobryidae. However, the boss and tooth disintegration are

not characters demonstrating relationship conclusiv ely,

as similar structures are also known from unrelated

groups (Malchus and Warén, 2005, for a recent discus-

sion) and microstructure details are still needed for other

Philobryidae.

ACKNOWLEDGMENTS

I thank Anders Warén for logistic and scientific support

during my stay at the Swedish Museum of Natural His-

tory, Stockholm, and Alan Beu (GNS Science, NZ), Di-

ego Zelaya (MNCN, La Plata, Argentina) and two anony-

mous reviewers for their constructive comments. Finan-

cial support through the European Commissions’

“Access to Research Infrastructure” (High Lat Re-

sources, HPRI-CT-2001-00125) and the Spanish Minis-

terio de Educacién y Ciencias and Generalitat de Cata-

lunya (Ramon y Cajal Research Contract) is highly ap-

preciated.

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THE NAUTILUS 120(1):15—-20, 2006

Descriptions of Otukaia crustulum new species (Gastropoda:

Trochoidea: Calliostomatidae) and Margarites huloti new species

(Gastropoda: Trochoidea: Trochidae) from a methane seep area

off Chile

Claude Vilvens

Rue de Hermalle, 113

B-4680 Oupeye

BELGIUM

[email protected]

Larrondo 1281

Coquimbo

CHILE

[email protected]

Javier Sellanes

Universidad Catoélica del Norte

Facultad de Ciencias del Mar

ABSTRACT

Two new species of Trochoidea are described from a methane

seep area off central Chile (~36°S). Otukaia crustulum new

species is compared with similar calliostomatid species such as

Otukaia kiheiziebisu (Otuka, 1939), O. eltanini Dell, 1990, and

Calliostoma jacquelinae McLean, 1970. The new species is

separated from these similar species by convex whorls bearing

a prominent abapical cord ee an almost invisible adapical

cord, and five spiral cords on the base. Margarites huloti new

species from the same locality is compared with similar mar-

garitine species, such as M. shinkai Okutani, Tsuchida and

Fusions, 1992, M. salmonea (Carpenter, 1864), and Solariella

tavernia Dall, 1919, and with calliostomatid species such as

Calliostoma nudiusculum (von Martens, 1881), C. magaldii

Caldini and Guimaraes-Prado, 1998 and C. keenae McLean,

1970. The new species differs mainly by the size of its shell,

elevated spire, rather thick spiral cords on the whorls, numer-

ous low spiral cords on the base and the narrow, funnel-shaped,

umbilicus.

INTRODUCTION

The deep-sea malacofauna of Chile is still scarcely

known. The bathyal (>200 m water depth) mollusks as-

sociated with the deep-sea shrimp fisheries off central

Chile (31 to 35°S) were described by McLean and An-

drade (1952). Among the 15 species observed, four be-

longed to the Trochoidea (sensu Bouchet et al., 2005),

two of them belonging to the genus Otukaia (O. chilena

Rehder, 1971) and O. delli (McLean and Andrade, 1982)

and two to the genus Bathybembix (B. macdonaldi (Dall,

1890) and B. huwmboldti Rehder, 1971).

The diversity of the Trochoidea from the coast of

northern Chile (15 to 31°S) was subsequently studied by

Véliz and Vasquez (2000). Their study cove ies the inter-

tidal to the slope area, and among the 11 species re-

ported, the same four species indicated by McLean and

Andrade (1982) were mentioned for the bathyal re gion.

Vilvens (2002) described Zetela alphonsi from 800 m

depth from off Chiloé, farther south (~40°S),

Forcelli (2000) cited 27 species of Trochoidea for the

Magellanic province, but almost always without consid-

ering their distributional range (except for Calliostoma

fonkii (Philippi, 1860) (Figures 8-9), clearly reported for

the Pacific side of the Magellanic province); some of

these species are in fact described from or reported for

the southwesterm Pacific (e.g., Calliostoma consimilis (1

A.Smith, 1S81))

In relation to the shallow-water malacofauna of Chile,

Calliostoma fonkii (Philippi, 1860) has been known from

moderately shallow-water north to Peru and the Galapa-

gos Islands (Keen, 1971). Several fossil species of the

genus Calliostoma have been found in Miocene sedi-

ments from the Arauco Peninsula (~38°S), none with

geological ranges extending to the Recent (Nielsen et al.,

2004) :

Recent dredge hauls taken 75 km NW off the Bay of

Concepcion have allowed the location of a previously

unknown habitat at bathyal depths along the Chilean

coast, a methane seep area (Sellanes et al., 2004) named

Concepcion Methane Seep Area or CMSA (Sellanes and

Krylova, 2005). At methane-seep areas, as well as in

other marine reducing environments, chemically re-

duced fluids are utilized as an energy source by free-

living and endosymbiotic chemosynthetic bacteria in the

synthesis of organic carbon (Paull et al., 1954). Further-

more, microbially-mediated oxidation of methane within

the sediments facilitates production of carbonates that

act as a cement, ultimately giving rise to reefs of carbon-

ate-cemented mud. A diverse community benefits both

from the food sources originating from chemosynthesis

and from the shelter and substrate provided by the reefs

(Sibuet and Olu, 1988),

Page 16

THE NAUTILUS, Vol. 120, No. 1

About 30 species of mollusks have been recorded at

the CMSA. Part of this assemblage is composed by sev-

eral recently described species of f chemosymbiotic bi-

valves (Holmes et al., 2005: Oliver and Sellanes, 2005:

Sellanes and Krylova, 2005), and non-chemosymbiotic

accompanying species (Sellanes, unpublished data).

Among gastropods, five species of Trochoidea have been

econded. including Bathybembix macdonaldi, Otukaia

chilena, Zetela alj shonsi, and the two unnamed species

proposed herein. Tn this paper we describe a species of

Otukaia and a species of Margarites that have been col-

lected at the CMSA.

MATERIALS AND METHODS

Material in the present study consists of specimens ob-

tained living (lv) from the dredgings of R/V VipaL

GORMAZ in various stations (stn) of the CMSA.

Abbreviations used for measurements are: H: shell

height; W: shell width; HA: aperture height; TW: num-

ber of teleoconch whorls. Spiral cords of teleoconch of

the shells are labelled as P1, P2, etc., for primary cords

(P1 is the most adapical) and S1, $2, etc., for secondary

cords (S1 is the most adapical).

Type specimens are deposited at Natural History Mu-

seum of Chile, Santiago (MNHNCL) and Institut royal

des Sciences naturelles de B Belgique, Bruxelles, Belgium

(IRSNB).

SYSTEMATICS

We follow herein the suprageneric classification of Mar-

shall (1995) and Bouchet et al. (2005) for Calliostoma,

Otukaia, and other related genera. Other authors, how-

ever, use the classification of Hickman and McLean

(1990) where Calliostomatinae is considered as a sub-

family of Trochidae.

Superfamily Trochoidea Rafinesque, 1S15

Family Calliostomatidae Thiele, 1924

Subfamily Calliostomatinae Thiele, 1924

Tribe Calliostomatini Thiele, 1924

Genus Otukaia Ikebe, 1942

Type Species: Calliostoma kiheiziebisu Otuka, 1939

(by original designation); Recent, off Japan.

Otukaia crustulum new species

(Figures 1-5)

Description: Shell of medium size for genus (height

up to 15.6 mim, width up to 14.4 mm), conical to weakly

coeloconoidal in shape; spire high, as broad as large,

height about 2.4 to 3.5 times higher than aperture; um-

bilicus closed in adult shell.

Protoconch damaged on all available specimens, about

150 jum in diameter, probably encompassing | whorl, too

eroded to state presence and shape of terminal varix,

Teleoconch of up to 6 moderately convex whorls, bear-

ing 3 spiral granular cords, adapical one the weakest, only

poorly visible on last whorls; prosocline ribs on first 3

whorls connecting beads on spiral cords and very thin,

axial lamellate threads between cords of the base. Suture

visible, impressed, not channeled. First whorl of teleo-

conch convex, sculptured by 2 primary cords; P2 appear-

ing immediately, granular; P1 appearing a quarter of

whorl later, subgr anular, weaker than P2: ; prosocline axial

ribs in the intervals between cords, conne cting beads of

cords; interval between ribs 1.5 times larger ian width

of ribs. On second whorl, beads of P2 becoming thicker

and sharp, isolated but connected by cord; beads of P1

axially elongated; P3 partly emerging ‘from suture, granu-

lar: interv al between axial ribs becoming 2 times larger

than width of ribs; area between P2 and P3 concave. On

third whorl, Pl is closer to suture, distance between

beads of about 1.5 times width of beads; P2 the strongest,

beads becoming axially elongated, distance beewecn

beads decrease to size of head B P3 clearly visible, weak-

est, with small sharp beads. On fourth whorl, P2 closer to

second abapical third of whorl, with beads less sharp,

axially very elongated, and closely packed; beads of Pl

becoming ‘weaker, se parated by interval 3 times larger

than bears: axial ribs becoming obsolete. On fifth whorl,

Pl becoming obsolete, almost disappearing; P2 thick,

beads reducing to — prosocline ribs. On last whorl, P1

virtually invisible; P2 very thick, producing carina; P3

much weaker, small beads. Aperture subquadrate;

interior of outer lip with lirae corresponding to external

cords, producing strong angle at the rim; inner lip

curved, projecting over agibilical area. Columella slightly

curved, slightly oblique, without tooth. Base weakly con-

vex, with 5 spiral cords; 2 innermost cords granular, in-

terspace between cords similar in size to cords: 3 outer-

most cords subgranular, distance between cords two

times larger than cords; very thin axial lamellate threads

between cords, stronger in umbilical area. Color of tel-

eoconch brownish-beige, tips of beads of spiral cords

lighter in color.

Type Locality: Central Chile, 36°21.91’ S, 73°43.21'

W, 843-728 m, South Pacific Ocean.

Type Material: Holotype MNHNCL (201649) (lv),

AGOR Vidal Gormaz, stn AGT 13, from type locality,

15.6 x 14.4 mm; Paratype IRSNB 30514 568 (lv), 10.4 x

11.0 mm.

Etymology: Of a cake (Latin), with reference to the

shape of the shell whorls, which invoke the image of a

iked cake. The genus-group name Otukaia was e rected

by Ikebe without dé finition of its etymology and without

reference to its gender. We assume that the name is

neutral.

Remarks: Absence of complete protoconchs in speci-

mens from the type series prevents a definitive allocation

of the new species in the Calliostomatidae. Provisional

placement in this family, however, is prompted by the

general shell shape, open umbilicus in the immature

specimen, and similarities in the axial sculpture of the

C. Vilvens and J. Sellanes, 2006

Figures 1-9. Ofwkaia and Calliostoma species. 1-5. Otukaia crustulum new species, Central Chile. 1-3. Holotype MNHNCL

201649, 15.6

Galapagc »s Islands, 12.0

14.4 mm. 4-5. Paratype IRSNB 30514, 10.4

early teleoconch whorls with other calliostomatids. Fur-

ther studies including more complete specimens may

reveal that the new species belongs in the Chilodontidae,

tribe Calliotropini.

Among Calliostomatidae, the genus Otukaia Ikebe,

1942. seems to be the most adequate to include the new

species Otukaia crustulum new species weakly re-

sembles O. kiheiziebisu (Otuka, 1939) from Japan

200-1000 m; Sasaki, 2000: pl.37, fig.108), but the me-

dian spiral cord P2 of this species is much thinner and

the base bears more cords. The new species may also be

compared to O. eltanini Dell, 1990, from the Pacific-

Antarctic Ridge (915-1153 m: Dell, 1990: fig.168); this

species, howe vel be ars { primary evenly distributed Spl-

11.0 mm. 6, 8. Calliostoma jacquelinae McLean, 1970, C.Vilvens coll

13.0 mm. 7, 9. Calliostoma fonkii (Philippi, 1860), C.Vilvens coll., northern Chile, 16.0

14.5 mm

ral cords on the last whorl, with 2 strong adapical cords

and 21 spiral cords on the base. Otukaia crustulum new

species is rather similar to Calliostoma jacquelinae

McLean, 1970, (Figures 6, 8) from the Galapagos Islands

(150-350 m). Both species have a prominent abapical

cord, but this latter species differs by having a strong

granular, subsutural primary cord Pl, about 8 low

smooth spiral cords between PL, and many more (about

30) narrow, smooth spiral cords on the base

Family Trochidae Rafinesque, 1S15

Subfamily Margaritinae Stoliczka, 1865

Tribe Margaritini Stoliczka, 1S6S

Genus Margarites Gray, 1S47

Page 18 THE NAUTILUS, Vol. 120, No. 1

Figures 10-19. Margarites species. 10-17. Margarites huloti new species, Central Chile; LO-12. Holotype MNHNCL 201650,

12.5 « 13.4 mm, 13. Paratype MNHNCL 201651, 9.5 « 11.5 mm; 14-17. Paratype MNIINCL 201651, features of the radula. 14.

reneral vic Seale bar = 100 wm. 15. Rachidian and lateral teeth. Scale bar = 10 jm; 16. Marginal teeth lateromarginal plate and

lateral teeth. Scale bar = 10 wm; 17. Details of lateromarginal plate. 18-19. Margarites shinkai Okutani, Tsuchida and Fujikura

1992. holot NSMT Mo 69635, Japan, 11.0 « 15.1 mm

C. Vilvens and J. Sellanes, 2006

Page 19

Table 1. Margarites huloti. Shells measurements in mm for the largest specimens (n = 4). Abbreviations are: H: shell height; W: shell

width; HA: aperture height; TW: number of teleoconch whorls.

Character TW H W HA H/W H/HA

Range 4.00-5.00 9.80-12.50 11.50-13.40 4.60-5.30 0.85—0.94 2.04-2.43

Mean 4.55 11.45 12.50 4.95 0.9] 2.3]

Standard deviation 0.38 1.10 0.82 0.27 0.04 0.16

Type species: Turbo helicinus Phipps, 1774, by origi-

nal designation; Recent, northern Atlantic Ocean.

Margarites huloti new species

(Figures 10-17)

Description: Shell of medium size for the genus

(height up to 12.5 mm, width up to 13.4 mm), cyrto-

conoidal in shape: spire moderately high, height about

0.9 times width and 2.0-2.4 times aperture he ight ( Table

1); umbilicus narrow and deep. Protoconch about 550

wm wide, about 1 whorl, smooth, with a thin terminal

varix. Teleoconch of up to almost 5 convex whorls, bear-

ing spiral cords; adapical cords granular, abapical cords

smooth. Suture visible, impressed, not channeled. First

whorl of teleoconch convex, sculptured by 6 smooth,

weak, low, very close, similar in size and sh: ape primary

cords (from P2 to P7): weak irregular growth lines. On

second whorl, cords becoming stronger except P2 still

weak: Pl appearing under suture, smooth: interval be-

tween cords of about half of width of cords. On third

whorl, S1 appearing, weak, weakly granular; cords evenly

spaced; distance between cords similar in size to cords:

area between cords concave, with prosocline thin growth

lines. On last whorls, $1 clearly granular, with axially

elongated beads; P2 becoming weakly granular, with

horizontally elongated beads; other cords smooth; dis-

tance between cords of about 1.5 times width of cords:

PS emerging from suture only on last whorl, close to P7:

S2 sometimes appearing at end of last whorl, thinner and

subgranular. Aperture oval; interior of outer lip with thin

lirae corresponding to the external cords; inner lip

weakly curved, projecting over the umbilicus. Columella

straight, slightly oblique, without tooth. Base nearly flat

or very weakly convex, with 20 to 25 smooth spiral cords:

three innermost broader than others; interspace between

cords smaller than cords, smooth or with very weak axial

threads. Umbilicus narrow, funnel shaped, « diameter

about 10% of shell diameter in largest specimens, with

crowded axial lamellae and no spiral cord within. Colour

of protoconch and teleoconch iridescent grey, last whorl

lighter: base whitish grey. Operculum horny, multispiral

with a short growing edge: Radula rhipidogloss: ite: for-

mula ca. 20 + (1) + 6 + 1 +64 (1) + ca. 20. Rachidian

and lateral teeth similar in size and shape, with large, oval

base and serrated overhanging cusps. Lateromarginal

plate present. Marginal teeth thin, with long shaft and

serrated cutting edges.

Type Locality: Central Chile, 36°21.91' S, 73°43.21'

W, 843-728 m. South Pacific Ocean.

Type Material: Holotype: MNHNCL 201650 (lv),

AGOR Vidal Gormaz, stn AGT 13, from type locality,

12.5 x 13.4 mm; paratypes: MNHNCL 201651 (5 ly),

IRSNB 30514 569 (1 lv), all from type locality.

Etymology: Named after Andre Hulot, Belgian hydro-

biologist, United Nations Development Programme, a

scientist who initiated the marine science efforts in 1956

at the University of Concepcién, Chile.

Remarks: Margarites huloti new species is superti-

cially similar to another methane see p species M. shinkai

Okutani, Tsuchida and Fujikura, 1992 (Figures 18-19)

from Japan. This latter species, however, has a more

depressed shell, whorls more convex, more numerous

and thinner spiral cords on whorls, radula with 8 lateral

teeth (in contrast to only 6 in the new species): in addi-

tion, the rachidian and lateral teeth in the Japanese spe-

cies have a more slender base (Okutani, Tsuchida and

Fujikura, 1992: figs. 15-16).

The new species seems rather similar to Margarites

salmonea (Carpenter, 1864) from southern California,

but the latter is much smaller for a similar number of

whorls, has less spiral cords on the base, and a broader

umbilicus. Margarites huloti new species may also be

compared to Solariella tavernia Dall, 1919, from the Ga-

lapagos Islands. This latter species, however, has a

smaller size for about the same number of whorls and

bears more convex whorls with thinner spiral cords. The

new species also resembles Calliostoma nudiusculwum

(von Martens, 1881) from the southwestern Atlantic, but

the latter species differs from the new species by having

a more depressed spire, less numerous spiral cords on

whorls, and lacks an open umbilicus. The new species

weakly resembles Calliostoma magaldii Caldini and

Guimaraes-Prado, 1998, from the southwestern Atlantic,

but shells of this latter species have no umbilicus, are

pinkish white, and have whorls and base bearing less

numerous and thicker spiral cords. Margarites huloti

new species may also be compared to C allicstonis keenae

McLean, 1970, from the central eastern Pacific, but this

latter species is slightly larger, has a more horizontally

elongated aperture, and thinner and more numerous

granular spiral cords on whorls.

ACKNOWLEDGMENTS

Our thanks to the officers and crew of R/V VIDAI

GorMaz, for their skilful assistance at sea. Funding for

ship time was provided by the Office of Naval Research

Page 20

THE NAUTILUS, Vol. 120, No. 1

(ONR). The Center for Oceanographic Research in the

Eastern South Pacific (COPAS) and the Research Direc-

tion of the University of Concepcion also provided partial

support. We are very especially grateful to J. L. Van

Goethem (Institut royal des Sciences naturelles de Bel-

gique, Brussels) for ‘his help with requisition of type

loans. We also would like to thank H. Saito (National

Museum of Science of Tokyo) for the loan of types from

his institution. Last but not least, we are indebted to

Maria Soledad Romero (Facultad de Ciencias del Mar,

Universidad Catélica del Norte), for help with the SEMs

of the radula of Margarites huloti.

LITERATURE CITED

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and A. Warén. 2005. Part 2. Working classification of the

Gastropoda. In: Bouchet, P. and J. P. Rocroi (eds.) Clas-

sification and nomenclator of gastropod families. Malaco-

logia 47: 240-397.

Dell, R. K. 1990. Antarctic Mollusca, with special reference to

the fauna of the Ross Sea. Royal Society of New Zealand

Bulletin 27: 1-311.

Forcelli, D. O. 2000, Moluscos Magallanicos: Guia de moluscos

de Patagonia y sur de Chile. Vazquez Mazzini Editores,

Buenos Aires, 200 pp.

Keen, A. M. 1971. Sea Shells of Tropical West America, 2nd

edition. Stanford University Press, Stanford, x + 1064 pp.

Hickman, C. 8. and J. H. McLean. 1990, Systematic revision

and suprageneric classification of trochacean gasteropods.

Natural History Museum of Los Angeles County, Science

Series, 35, vi + 169 pp.

Holmes, A., P. €

Lucinoma ( Biv alvia: Lucinoidea) from a methane gas seep

off the southwest coast of Chile. Journal of ¢ ‘onchology 35:

673-681

Marshall, B. A. 1995. Calliostomatidae from New Caledonia,

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pagnes MUSORSTOM, Volume 14, Mémoires du

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THE NAUTILUS 120(1):21-24, 2006

Page 2]

A new land snail of the genus Humboldtiana (Gastropoda:

Pulmonata: Mumbolddanidae) from Nuevo Leon, Mexico

Fred G. Thompson

Florida Museum of Natural History

University of Florida, Box 117800

Gainesville, FL 32611-7800 USA

[email protected]

ABSTRACT

Humboldtiana iversoni new species is described from southern

Nuevo Leén, Mexico. The species is most similar to H. mon-

tezuma Pilsbry, 1940, because of its reproductive system mor-

phology and shell sculpture. The new species differs from H.

montezuma by its smaller size, its color pattern and aspects of

the structures of its reproductive system. It is unique within the

genus because of its bulbous penis and its short, broadly conical

vagina. Because of features of the dart apparatus in the female

reproductive system Humboldtiana inferior Pilsbry, 1948, is

recognized as a distinct species from H. montezuma, of which

it was considered a subspecies formerly.

INTRODUCTION

Humboldtiana is a genus of large helicoid land snails

(Gastropoda, Pulmonata, Helicoidea, Humboldtianidae).

The genus is distributed from Texas and Chihuahua

south to areas near Mexico City and central-western Ve-

racruz. feel) fifty species are recognized

(Thompson, in press), and it is apparent that many more

wait to be discovered. One such novelty was collected by

John B. Iverson many years ago during he ‘rpetological

investigations in northeastern Mexico. Repository insti-

tutions are: UF: Florida Museum of Natural History,

University of Florida; ITCV: Instituto Tecnologico de

Ciudad Victoria, Tamaulipas, Mexico.

SYSTEMATICS

Family Humboldtianidae Pilsbry, 1939

Genus Humboldtiana von thering, 1892

Humboldtiana iversoni new species

Description: The shell (Figures 1-4) is small to me-

dium for the genus, helicoid, up to about 35 mm wide,

about 0.S5-1.00 times as high as wide. The color pattern

consists of three narrow brown bands (Figures 1, 3). The

middle band is the most conspicuous. The bands are

partially interrupted by white riblets on a tan or light-

brown background color. The internal surface of aper-

ture is tinged light tan in fresh specimens. The bands are

barely distinguishable within the aperture. The suture is

shallow, and ‘dexeeane s abruptly to the aperture from the

middle band to the lower band along the last eighth

whorl. The whorls are weakly shouldere d and nearly uni-

formly rounded. Adult shells have 4.0-4.6 whorls. The

embryonic shell consists of 1.4-1.6 whorls. The first em-

bryonic whorl is smooth, flat-topped, and 2.25 mm wide

pe rpendicular to the initial suture. The following half

whorl is covered with numerous very fine e lliptical gran-

ules and has faint radial striations below the pe riphery.

The post-embryonic whorls are sculptured with numer-

ous close grow th wrinkles and dense, fine, e longate gran-

ules that are aligned transversely (Figure 4). On the last

1.5 whorls the wrinkles become well-defined riblets that

are about as wide as their interspaces. The riblets are

continuous into the umbilicus. Transverse rows of gran-

ules are superimposed on and between the riblets. The

aperture is nearly round in frontal view and is 0.76-0.82

times as wide as high in one view. The plane of the

aperture lies at an angle of 37-43° to shell axis. The

posterior corner is rather wide ly separated from the col-

umellar margin. The parietal ‘callus consists of a thin

transparent glaze. The peristome is a and blunt-

edged. The basal-columellar edge is slightly reflected.

The columellar lip is narrowly réfle cted to partially cover

the umbilicus (Figure 2). Shell measurements in mm

based on the holotype and the three paratypes are given

in Table 1

Anatomy (Figures 5-6): (Morphological terminology

follows Thompson and Brewer, 2000.) The holotype is

the only specimen that was available for study. It was

preservec ed in the field in 70% ethanol, and was tightly

contracted into the shell. The head-foot is dark gray. The

mantle over the lung is uniform lighter gray. Reproduc-

tive system (Figure 5): The genité i atrium is very short.

The penis has a shove narrow neck, and then it rapidly

expands into a wide, bulbous orb that is wider than high;

length of penis 4.5 mm. The penis wall is very thin

Internally the wall has several longitudinal gl: indular

folds in the neck. The penis has a very large verge that

fills the upper half of the cavity and e mds in four heavy

age 22

Page 22

THE NAUTILUS, Vol. 120, No. 1

Figures 1-4. Humboldtiana iversoni new species. Shells. 1-2. Holotype, UF 103588. 3-4. Paratypes, UF 367594. 3. Paratype A.

4. Paratype B. Scale bar for Figures 1-3 = 10 mm. Figure 4 is enlarged 25% in relation to Figures 1-3.

lobes (Figure 6). The penis retractor muscle (pr) is short

and stout; length 3.5 mm. It originates on the center of

the mantle cavity about 1/8 of a whoel behind the mantle

collar and inserts on the base of the epiphallus (epi)

juxtaposed to the penis. It does not form a sheath around

the epiphallus. The epiphallus is 11.5 mm long. It is

moderately slender and uniformly wide throughout its

length, bemg about the same diameter as the neck of the

penis. It is lined internally with four longitudinal folds.

The flagellum (flg) is 13 mm long. It is ‘slightly longer

than the epiphallus, is moderately stout, sanidl bears four

longitudinal folds internally. The vagina (vg) is short and

stout with a broad conical base: total length of vagina 7.5

mm. The vagina bears four dart sacs (ds) of equal size.

Dart bulbs are not evident externally, The dart glands

(dgl) form around the vagina a robust ring that is juxta-

posed to the dart sacs. The f free vagina (vet) i is robust and

is about 2.5 mm long. The sper miathecal duct (sptd) is 27

min long. It bears a caecum (cae) 16 mm above it base.

The caecum is 9 mm long. The spermatheca is oval in

shape and is relatively small for the genus. Measure-

ments could not be made of the uterus (utr) or the

albumen gland because of their state of preservation.

Type Material: Holotype, UF 130588; Paratypes, UF

367594 (2), ITCV (1); same data as the holotype; all

collected by John B. Iverson, 18 June 1978, at type lo-

cality.

Type Locality (Figure 7): Nuevo Leon; Highway 65

at km post 116, Las Norias (24° 11.07 N, 99°53.0° W),3

kim south of Anteojitos; 1SO00 m altitude. Las Norias is

about 50 km SE of Galeana on the highway to La Es-

condida and Doctor Arroyo. Specimens were found un-

der limestone blocks on an open southeast facing grassy

hillside with nearby scattered oaks (Quercus sp.) and

clusters of Agave sp.

Table 1. Hiumboldtiana iversoni new species. Shell measurements in min of the holotype and three paratypes. H: shell height; W:

shell width; AW: aperture width; AH: aperture height; AA: angle of aperture to shell axis.

H Ww AW Al Whorls H/W AW/W AH/H AW/AH AA

Holotype 28.6 28.5 16.0 20.6 4.4 1.00 0.56 0.72 0.78 37°

UF 27.3 29.2 16.4 21,7 4.3 0.93 0.56 0.79 0.76 39°

UF 24.4 28.8 15.] 18.5 4.0 0.S5 0.52 0.76 0.82 3°

ITCV 30.6 35.6 17.5 22.06 4.6 0.9] 0.50 0.75 0.78 40°

F. G. Thompson, 2006

Page 23

5 a

enis

Pp 10 mm

ee eee) ee ee ee |

Figures 5-6. Humboldtiana iversoni new species. 5. Reproductive system. 6. Interior of penis.

Distribution: The species is known only from the type

locality.

Etymology: This species is named for John B. Iverson,

Earlham College, Richmond, Indiana, in acknowledge-

ment for this and other novel species of mollusks col-

lected by him in Mexico while conducting herpetological

explorations.

Remarks: =Humboldtiana iversoni belongs in the sub-

genus Humboldtiana, or at least, what is thought to be

the subgenus Humboldtiana. The identity of he type

species, Helix humboldtiana Pfeiffer, 1857, remains

equivocal, ae its type locality and range are unknown

(Pilsbry, 1927). It was described from “Mexico” without

a specific iooliey Fischer (1899) described and illus-

trated the anatomy of a specimen from Guanajuato that

he identified as H. humboldtiana. but he did not describe

or illustrate the shell, nor is it clear whether he meant the

City of Guanajuato or the State of Guanajuato. For these

reasons Pilsbry (1927) expressed reservation concerning

the identity of Fischer's specimen. Notwithstanding the

limitations concerning the identity of the type species,

the subgenus is tentatively characterized as follows, be-

cause all other species described in the 19'* Century and

all species subsequently described from central and

southern Mexico are known anatomically, and there is

little reason to doubt that H. humboldtiana departs sig-

nificantly from a common ground plan. This includes

Fischer's description. oa species have in common the

following characteristics: (1) The spermathecal duct has

an appendix: (2) There are four dart sacs on the vagina;

(3) Two dart bulbs are imbedded in the vaginal w. all at

the base of each sac and are not visible externally; (4) The

four dart glands coalesce immediately above the dart sacs

to form a ring around the vagina; and (5) The penis

contains a verge internally, which is a papilla-like exten-

sion of the epiphallus into the penis chamber.

Humboldtiana iversoni belongs to a small group of

species that includes H. montezuma Pilsbry, 1940, and H.

inferior Pilsbry, 1948 (formerly Humboldtiana monte-

zuma inferior Pilsbry, 1948). The group is found in

southeastern Nuevo Le6én. Humboldtiana montezuma is

found at 3080 m altitude at the summit of El Infiernillo,

a mountain near Pablillo, south of Galeana; and H. infe-

rior is found lower at about 1850 m altitude, near Pab-

lillo. The species have in common coarse postembryonic

Figure 7. Type locality of Humboldtiana iversoni new spe-

cies at Las Lorias, Nuevo Le6én, Mexico.

Wien Ol

Page 24

THE NAUTILUS, Vol. 120, No. 1

sculpture of close thread-riblets and minute beaded

granules that are aligned on or between the riblets. The

color pattern consists of light-colored ribs on a light

brown or yellowish background. Spiral bands are absent

in adult shell, or when present they are weakly defined.

The female reproductive system has four dart sacs on the

vagina, and the dart glands form a ring around the vagina

just above the dart sacs.

Humboldtiana iversoni is unique within Humboldtiana

because of the structure of the penis with its very large

bulbous verge. Also, it is unusual because of the short,

broadly conical lower vagina. Superficially the pen’ re-

sembles that of H. tescola Thompson, 1967. In the latter

species the inner wall of the penis bears heavy glandular

folds that surround a short and relatively slender verge,

and the penis retractor muscle inserts on the epiphallus

(Thompson, 1967). Humboldtiana iversoni differs from

other members of the H. montezuma species-group by

having a short, stout flagellum that is only slightly longer

than the epiphallus, and by having the penis retractor

muscle insert on the base of the ~epiphallus. Both H

montezuma and H. inferior have a long slender flagellum

that is 1.3-1.6 times as long as the epiphallus, and the

penis retractor inserts higher on the epiphallus (Pilsbry,

1948). Humboldtiana iversoni is like H. montezuma by

having four equal-sized dart sacs on the vagina. In H.

inferior dart sacs ds, and ds. are reduced in size com-

pared to dsj. Humboldtiana montezuma lacks bands,

while H. inferior has three bands. In both H. montezuma

and H. inferior the internal surface of the aperture is

brown, in contrast to light tan-tinged aperture of H. iver-

soni.

Humboldtiana inferior had been regarded as a subspe-

cies of H. montezuma. Pilsbry (1948) states that the dart

sacs of H. inferior are sub-equal in size, and his Fig, 1B

shows that at least ds, and ds, are conspicuously sqnailes

than is ds,. The reduced size of dart sacs ds, and dsy is

a sufficient basis for recognizing Humboldtiana inferior

as a separate species (see Thompson and Brewer, 2000).

ACKNOWLEDGMENTS

The anatomical drawings in Figures 5-6 were produced

by Susan Trammell, Gainesville, Florida. Her excellent

illustrations greatly enhance this paper. I am grateful to

two anonymous reviewers whose comments improved

this paper.

LITERATURE CITED

Fischer, H. 1899. Note sur Helix hiumboldtiana Valenciennes

avec queques remarques sur le sous-genre Lysinoe et sur

la section waa Journal de Conchyliologie 47; 297—

304, figs. I=

Pilsbry, H. A. 1997, The structure and affinities of Humboldti-

ana and related helicid genera of México and Texas. Pro-

ceedings of the Academy of Natural Science of Philadel-

phia 79: 165-192, pls. 11-14.

Pilsbry, H. A. 1940. Two new species of Mexican Humboldti-

ana. The Nautilus 53: 140-141.

Pilsbry, H. A. 1948. Island Mollusca of northern Mexico. I. The

genera Humboldtiana, Sonorella, Oreohelix and Ashmu-

nella. Proceedings of the Academy of Natural Sciences of

Philadelphia 100: 185-203; pls. 12-14.

Solem, A. 1954, Notes on Mexican mollusks. I Durango,

Coahuila and Tamaulipas, with descriptions of two new

Humboldtiana. The Nautilus 68: 3-10; pl. 1

Thompson, F. G. 1967. A new helicid snail from Zacatecas,

México. Nautilus. $1(1): 22-27; Figures 144.

Thompson, F. G. and G. P. Brewer. 2000. Landsnails of the

genus Humboldtiana from northern Mexico (GAS-

TROPODA, PULMONATA, HELICOIDEA, HUM-

BOLDTIANIDAE),. Bulletin of the Florida Museum of

Natural History 43: 49-77.

Thompson, F. G. In press. Some land snails of the genus Hum-

boldtiana from Chihuahua and western Texas. Bulletin of

the Florida Museum of Natural History.

THE NAUTILUS 120(1):25-29, 2006

Page 5

Two new land snails of the genus Humboldtiana (Gastropoda:

Pulmonata: Humboldtianidae) from Chihuahua, Mexico

Fred G. Thompson

Florida Museum of Natural History

University of ne Box 117800

Gainesville, FL 32611-7800 USA

Eeicai ile du.

Omar Mejia

Mexico, DF

MEXICO

Escuela Nacional de Ciencias Biolégicas

Instituto Politecnico Nacional,

[email protected].

ABSTRACT

Humboldtiana corruga new species and Humboldtiana sylva-

nia new species are described from Chihuahua. They are char-

acterized by anatomical features as well as shell morphology.

INTRODUCTION

Snails of the genus Humboldtiana comprise some of the

larger land snails in Mexico. In most cases the species

occur in sparsely populated colonies, and they have in-

sular distribution patterns. Uniform distributions over

areas larger 1 km? are rare in what appear to be appro-

priate habitats. Instead, species occur in isolated colo-

nies, with local endemism being the rule. This study

describes two such species. Their descriptions are

prompted by the need to round out phylogenetic studies

(Mejia. in prep.) We use the following abbreviations for

shell measurements: H: height; W: wadthe AH: aperture

height; AW: aperture width. Repository institutions are:

ITCY, Instituto Tecnolégico de Ciudad Victoria,

Tamaulipas, Mexico; UF, Florida Museum of Natural

History, University of Florida, Gainesville, Florida.

SYSTEMATICS

Family Humboldtianidae Pilsbry, 1939

Genus Humboldtiana von Ihering, 1892

Humboldtiana corruga new species

Diagnosis: This is a moderately large species that has

smooth embryonic whorls. The g lobose shell is about as

high as wide. It is sculptured with rugose growth wrinkles

on the postembryonic shell, with dense granular sculp-

ture aligned with the growth wrinkles. The color pattern

consists of three narrow blackish bands of equal size on

a dark brown ground color. The whorls are inflated and

tend to take on a square appearance. The last whorl has

a distinct s teat and has a slightly channeled suture.

The edge of the peristome is blunt and not reflected.

Description: The shell (Figures 1-5, Table 1) is me-

dium in size, about 32 mm wide, globose in shape, about

1.02 times as high as wide. The color is lusterless dark

brown with lighter brown streaks along the growth lines,

and with three nearly equally narrow black vee The

bands are well defined, although they tend to be dis-

rupted by transverse streaks on ‘he lower half of the body

whorl. The aperture is lighter brown and banded inter-

nally. The shell has up to 4.0 whorls. The 1.5 embryonic

whorls protrude conspicuously above the following

whorls. The first embryonic whorl is 5.8 mm wide trans-

verse to the initial suture. The postembryonic whorls are

inflated with a noticeable shoulder and a channeled su-

ture which tends to give the whorls a squared appear-

ance. The last half- sani gradually descends to the ap-

erture, The peristome inserts on the lower edge of the

lower band. The embryonic whorls are smooth ( Figures

3, 5). The postembryonic whorls are sculptured “with

coarse growth wrinkles and striations, which are continu-

ous to fhe peristome and into the umbilicus. Numerous

minute granules are superimposed on the growth

wrinkles. The granular sculpture extends from the suture

to the base, but it does not continue into the umbilicus.

The aperture is wrinkled internally beneath the outer

sculpture. The aperture is 1.3 times as high as wide and

is about 0.73 times the height of the shell. It is prosocline

at an angle of 18° to the shell axis (Figure 2). The peris-

tome is blunt- edged, and is not reflected along the up-

per, outer, and hacal lip, but it is broadly refle cted along

the columellar lip to partially cover the umbilical area

(Figure 4).

Anatomy (Figures 11, 12): The anatomy of the holo-

type is ¢ lescribed. The head-foot is gray above and on the

sides bordering the sole. A lighter gray zone extends on

the sides from the snout posterior to the mantle collar.

The sole is very light gray. The mantle collar is light gray.

The outer wall of the mantle cavity is light gray and is

reticulated with narrow black lines.

The genital atrium (gen at) is slender, and is about a

third the length of the penis (Figure 11). The penis is 14

mm long and is bulbous with a slight constriction near

Page 26

THE NAUTILUS, Vol. 120, No. 1

Figures 1-10. New species of Humboldtiana. Shells. 1-5. Humboldtiana corruga new species. Holotype, UF 358872. 6-10.

Humboldtiana sylvania new species. Holotype, UF 353714.

the apex at the insertion of the penis retractor muscle

(pr). The penis has a verge that extends about half the

length of the cavity. The verge is surrounded by a heavy

pendulant curtain of olandular tissue (Figure 12). The

lower wall of the penis bears a few small longitudinal

folds below the curtain. The penis retractor aneccle (pr)

is 6 mm long, and is relatively short and moderately

stout. It originated on the inner wall of the ling slightly

behind the Taidelle of the mantle collar, and inserts on

the apex of the penis where it forms a narrow sheath

around the base of the epiphallus. The epiphallus (epi) is

slender, and is slightly longer than the penis. It is lined

internally with four longitudinal folds. The slender fla-

cellum (flg) is about as long as the combined length of

the penis + epiphallus. The. vagina is about 16 mm long.

The lower vagina (vag,) has a long slender neck, and

bears four dart sacs of equal size (ds,, dsy, ete)) each

with a pair of dart bulbs (db) distinctly protruding at its

base. The bulbs are embedded in the wall of the vagina

and form conspicuous bulges in the wall. The dart glands

(dgl) are widely sep: Aenea from the dart sacs. The

middle vagina (vag,,) is about as long as the lower va-

gina, and is considerably stouter. The free vagina (vag,)

is very short, and is barely visible above the doit glands.

The spermatheca (spt) is small and globular. The sper-

mathecal duct (sptd) is very long; the combined length

of the spermatheca + duct is 65 mm. The duct bear a

caecum (eae) at about a third of the distance below the

spermatheca. The length of the uterus-prostate is about

40 min.

Type Material: Holotype, UF 358872, collected by

Omar Mejia, 25 August 2003; Paratype, ITCV (1 shell);

Table 1. Humboldtiana corruga new species. Measurements

of the holotype and the paratype. Measurements of the

paratype are incomplete because it has a fractured lip and apex.

Abbreviations used are: H: height; W: width; AH: aperture

height; AW: aperture width.

ul W AW AH Whorls

Holotype 31.5 31.0 17.6 23.0 3.8

Paratype _ 32.5 18.5 — 4.0

F. G. Thompson and O. Mejia, 2006 Page 27

Figures 11-14. New species of Humboldtiana. Reproductive anatomy. 11-12. Humboldtiana corruga new species. 11. Repro-

ductive systern. 12. Interior of penis. Abbreviations: agl: albumen gland; cae: caecum; db: dart bulb; dg: dart gland; ds: dart sac

epi: epiphallus: flg: flagellum; gen at: genital atrium; pr: penis retractor; pro: prostate; spt: spermatheca; sptd: spermathecal duct;

utr: uterus: vag: vagina: vd: vas deferens. 13-14. Humboldtiana sylvania new species. 13. Reproductive system. 14. Interior of penis

Page 28

THE NAUTILUS, Vol. 120, No. 1

both from the type locality. Measurements are given in

Table 1.

Type Locality: Chihuahua, 0.8 km south and 0.3 km

west of Norogachi, Chihuahua (27°15.9° N, 107° 7.8° W),

2280 m altitude, open Pinus-Quercus woodland.

Distribution: Known definitely only from the type lo-

cality.

Etymology: The species name corruga derives from

the Latin, co, meaning together or with, aid ruga, mean-

ing a wrinkle or fold. The name alludes to the coarse

wrinkled sculpture on the shell.

Remarks: Humboldtiana corruga differs from other

known species by its distinct anatomical features. Hum-

boldtiana corruga is unique within the genus because of

the following Combination of saatornt al characters. It

has a long genital atrium. The penis has a moderately

long verge, which is surrounded by a curtain of glandular

tissue. The lower vagina has a long slender neuk The

middle vagina is long. and widely separates the dart sacs

from the dart glands. The vagina bears four dart sacs of

equal size, aah, of which is hon dered by two dart bulbs.

The spermathecal duct has a relatively long caecum that

is located at about two-thirds of the distance from the

base.

Humboldtiana corruga superficially is similar to an-

other undescribed species from near San Ignacio

Arareco, southeast of Creel, Chihuahua in that the shells

are similar in size, and have similar color patterns

(Thompson, in press). In H. corruga the umbilical per-

foration is more conspicuous, the aperture is proportion-

ally higher, the suture is more a impressed and is

channeled al ong the body whorl, the bands are narrower

but better defined, and the bands are equal in width.

A very closely related form, which we tentatively iden-

tify as dete corruga, comes from Baqueachi,

Chihuahua (27°26.3’ N, 107°30.3 W), 1940 m altitude

(UF 359518). Baan is about 8 km west of Noroga-

chi. Our only available specimen has an immature shell.

Its reproductiv e system is virtually identical to that of H.

corruga, and its shell has similar ‘sculpture.

Humboldtiana sylvania new species

Diagnosis: This species is distinguished by its large

size, globose shape, and rotund ‘hols with a dictinet

shoulder. The color is straw yellow with three well-

defined black bands. The embryonic whorls are smooth.

The postembryonic sculpture consists of strong incre-

mental striations and ge Granular sculpture is ab-

sent over the surface of the shell.

Description: The shell (Figures 6-10) is large, up to

36 mm wide. It is sgntly depressed-globose, 0.88 times

as high as wide, and is thin shelled, and is shiny. The

color pattern is straw yellow with three distinct uninter-

rupte a black bands with well-defined straight edges. The

upper two bands are about equal in ade, The lower

band is narrower. The interior of the aperture is tan and

shows the external bands. The umbilical perforation is

narrowly rimate due to the reflected columellar margin

of the peristome (Figure 9). The shell has 4.0 rotund

whorls that are noticeably shouldered with a deeply im-

pressed suture. The body whorl descends to the aperture

along the last quarter turn. The 1.5 embryonic whorls are

shisoth (Figures 8, 10). The first embryonic whorl is 5.1

mm wide transverse to the initial suture. The post-

embryonic whorls are sculptured with relatively strong

incremental striations and wrinkle which extend undi-

minished from the suture to the umbilicus. Granular

sculpture is absent on all parts of the shell. The rotund

aperture is 0.80 times as high as wide and is 0.81 times

the height of the shell. The upper lip inserts between the

middle and the lower bands. The aperture is prosocline,

lying at an angle of 33° to the shell axis. The peristome is

blunt, nomouly but distinctly reflected along the upper,

outer and basal lips, and is broadly petlecied over the

umbilical area.

Anatomy (Figures 13, 14): Only the holotype was

available for dissection. The head-foot is very dark gray,

almost black. The sole is a slightly lighter shade of gray.

The mantle is light gray and is miofiled with dark gray

spots.

The genital atrium (gen at) is very short, almost non-

existent (Figure 13). The penis is bulbous with a slight

constriction below the apex. The interior of the penis has

a moderately slender verge that extends about half the

length of the chamber and is surrounded by five heavy

glandular folds (Figure 14). The neck of the penis is lined

internally with 6-7 low narrow longitudinal folds. The

penis retractor muscle (pr) is very short and stout. It

originates on the inner lung wall immediately behind the

toidele of the mantle collar, inserts on the apex of the

penis and forming a short sheath around the base of

epiphallus. The epiphallus (epi) is relatively slender for

the genus and tapers distally. It is 17 mm long and is

almost twice the length of ‘the penis. The interior of

gases is lined ith four longitudinal folds. The fla-

gellum (flg) is moderately long, ond is about as long as

the combined length of the penis + epiphallus. The . in-

terior of the flagellum is lined with four longitudinal

folds. The vagina is 30 mm long. The lower vagina (vagy)

is short and tapers below to a narrow neck. It bears four

dart sacs of equal size (ds,, dsy, ete.), each of which

bears a pair of dart bulbs (db) along its base. The dart

glands (dgl) form a well-dev eloped lobed ring around

fie y vagina. They are widely separated from the ‘dati sacs

by the middle vagina (vag,,), which is slightly longer

than the lower vagina. The free vagina (vagy) is very

short. The spermathec cal duct (sptd) is very long and

lacks a caecum. The spermatheca (spt) is small and ovate

in shape. The combined length of the spermatheca +

duct is 65 mm. The length of the uterus-prostate (utr-

pro) is 43 min.

714, collected by Omar Mejia, 31

32.0 mm: width: 36.2 mm: aper-

Holotype: UF |

August, 2003. He oi

F. G. Thompson and O. Mejfa, 2006

Page 29

ture height: 26.0 mm; aperture width: 20.9 mm; whorls:

4.0.

Type Locality: Chihuahua, Corareachi, 4.4 km north,

0.4 km west of Baqueachi (27°28.45° N, 107°30.93° W);

2000 m altitude. Corareachi is a very small village, which

is unnamed on the INEGI topographic map series 1:

50,000 (G13A82). The type locality is in a Pinus-Quercus

forest in a grassy glen.

Distribution: Known only from the type locality.

Etymology: The species name sylvania derives from

the Latin word for woods or forest.

Remarks: Shell and anatomical features in Humbold-

tiana sylvania are similar to those of an unnamed species

from near Rancho Blanco, NNE of San Juanito, Chihua-

hua (Thompson, in press). Important shell and anatomi-

cal traits that the two species have in common separate

them from other Humboldtiana. They share the follow-

ing shell traits. The embryonic whorls are smooth. The

poste embryonic whorls are sculptured with coarse incre-

mental striations and wrinkle. Granular sc ulpture is ab-

sent over the entire surface of the shell. The color pat-

tern consists of three black bands on a lighter back-

ground.

In addition, they share the following anatomical traits.

The outer wall of the lung is light gray and is mottled

with numerous small darker. gray spots. The genital

atrium is very short and alinost non-existent. The bul-

bous penis has a moderately slender verge that is sur-

rounded in the lower half by heavy longitudinal glandular

folds. The epiphallus is relatively sle a The flagellum

is about as long as the combined length of the penis +

epiphallus. The vagina bears four dave sacs of equal-size

that are widely separated from the dart glands. The sper-

mathecal duct lacks a caecum.

Aside from size, the anatomies of the two species are

very similar. The shells are distinguished by size and its

color. The unnamed species from near Rancho Blanco

has a shell that is up to 27 mm wide. Its ground color is

light diaphanous brown with three dark brown bands

that are interrupted by irregularly spaced alternating

dark brown transverse bars and blotches. This contrasts

strongly with color of Humboldtiana sylvania, in which

the straw yellow shell has with three distinct black bands

that have discrete edges.

ACKNOWLEDGMENTS

The anatomical drawings comprising Figures 11-14 were

rendered by Michael Falck, Gainesville, Florida.

LITERATURE CITED

Burch, J. B. and F. G. Thompson. 1957. Three new Mexican

land snails of the genus Humboldtiana. Occasional Papers

of the Museum of Zoology, University of Michigan, 590:

1-11.

Thompson, F. G. In press. Some landsnails of the genus Hiim-

boldtiana from Chihuahua and western Texas. Bulletin of

the Florida Museum of Natural History.

THE NAUTILUS 120(1):30-33, 2006

Page 30

Mollusks in a Holocene lake-sediment core from the Arctic

Foothills of northern Alaska

W. Wyatt Oswald!

College of Forest Resources and

Quaternary Research Center

University of Washington

Seattle, WA 98195 USA

INTRODUCTION

Mollusk remains preserved in lake sediments have been

analyzed in a variety of geographical regions and envi-

ronmental se ttings. These faunal records provide insights

into a range of issues, including (1) variations in mo Tusk

community composition and Cine (2) the current

aie el of freshwater gastropods and bivalves, and

) past changes in the environment (e.g., Watts and

anebe 1968; Harris and Pip, 1973; Hu et al., 1996;

Miller et al., 2000; Welter-Shultes and Richling, 2000;

Jensen et al., 2002; Mouthon and Magny, 2004). This

paper presents the first record of long-term changes in

the mollusk community of a lake from northern Alaska.

STUDY AREA

Red Green Lake (informal name; 68°39.2' N, 149°24.6'

W) is located north of boreal treeline in the central Arctic

Foothills of northern Alaska. The climate of this area is

cold, with average January and July ates ealaoat of

—22°C and 11°C, respectively; it receives ~325 mm mean

annual precipitation, most of which occurs during the

summer months (Zhang et al., 1996). The lake is ~2 ha in

area, with a maximum depth of 3.2 m, and pH of 8.2. Its

~50 ha watershed is dominated by prostrate-shrub tun-

dra plant communities (Walker et al., 1994, 1998). The

lake has a single outlet stream that flows west into the

Ithillik River.

MATERIALS AND METHODS

A 285 cm sediment core was recovered from Red Green

Lake in May 1997. Details of the coring operation and

subsequent laboratory analyses of the core’s organic con-

tent, chronology, and pe iynological record are described

elsewhere (Oswald a al., 2003). After the preliminary

sampling, the core was sectioned at 1 cm intervals and

' Present address: Harvard University, Harvard Forest, 324

North Main Street, Petersham, MA 01366, USA; woswald@

fas.harvard.edu

washed through a 500 wm screen. Mollusk remains were

identified by T. J. Frest using a modern specimen col-

lection and a cae of solerenwes (Burch, 1972, 1975

1989; Clarke, 1973, 1981; Baxter, 1957; Turgeon, 1998).

The mollusk data are expressed as the minimum number

of individuals (MNT) ) per 20 cm”. For bivalves, single right

and left valves were counted separately, and the lar ger of

those values was added to the number of paired v ail es to

arrive at the MNI. For gastropods, only those specimens

with intact apical whorls were included in the MNI value,

as adults could yield a number of large fragments per

individual. The specimens are deposited at the Quater-

nary Research Center, University of Washington, Seattle.

RESULTS

Six mollusk species (four bivalves, two gastropods) were

present in a 79 cm interval of the core dating to ~SO00—

4000 calibrated ''C years before present (cal yr BP) (Fig-

ure 1). The fauna for this interval included Pisidium lil-

jeborgi, Pisidium nitidum, Pisidium rotundatum,

Sphaerium nitidum, Lymnaea atkaensis, and Valvata sin-

cera helicoidea (Table 1). Mollusk diversity was low at

any given level in this interval, with an average of 1.91

species per sample, and a range of 1-4 species per

sample. P. lilieborgi was the most abundant bivalve spe-

cies, occurring in 46 samples between $300 and 4100 cal

yr BP, and reaching a peak abundance of 15 MNI at 6500

cal yr BP. The other Pisidium species were less common:

P. nitidum was found in two samples (5900 and 5400 cal

- BP), and P. rotundatuwm was found in five samples

ie 7100 and 6400 cal yr BP. Sphaerium nitidum

occurred in 13 samples between 7900 and 4600 cal yr

BP, whereas L. atkaensis was encountered in 18 samples

between 7900 and 5200 cal yr BP. V. sincera helicoidea

was the most abundant gastropod species, appearing in

23 samples between S000 and 4100 cal yr BP, with high-

est MNI values at SO00-7700 and 6800-6100 cal yr BP.

DISCUSSION

The faunal community present in the mid-Holocene in-

terval of the sediment core suggests that Red Green was

W.W. Oswald, 2006

Page 3]

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8 5 :

® AS) 3

= ‘< Oo

§ § §

ES) cS) 3

2) G 7)

a a a

1000

2000

3000

4 4000

[@))

= 5000

6000

7000

8000

9000

10 20 4 8

Minimum number of individuals

Figure 1.

Sphaerium nitidum

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ro)

(7a) a.

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5 .

pee

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4 4 10 20 4

Number of species

Changes over time in the abundance of mollusk species in a sediment core from Red Green Lake, northern Alaska. Y

axis is calibrated C years before present (cal age BP), Abundance value (x axis) of the mollusks is the minimum number of

individuals per 20 cm?

a shallow, oligotrophic lake between ~8000 and 4000 cal

yr BP. In particular, P. nitidum and P. rotundatum are

indicative of shallow lake depth, as they are common in

water <1 m deep (Clarke, 1981; G.L. Mackie pers.

comm.). S. nitidum, P. liljeborgi, and L. atkaensis can

also occur in shallow water, and they are most common

where oligotrophic conditions exist (Clarke, 1979, 1981;

G.L. Mackie pers. comm.). Valvata sincera helicoidea

indicates that ac juatic vegetation was present, as it is typi-

cally found with aquatic plants (Clarke, 1973). The in-

terpretation of these species as indicators of shallow wa-

ter depth is consistent with the findings of paleoecologi-

cal studies in south-central Alaska (Hu et al., 1996) and

in the northwestern corner of Canada’s Northwest Ter-

ritories (Delorme et al., 1977).

The decline in mollusk abundance between ~5500 and

4000 cal yr BP suggests that the lake became deeper

during that interval. That interpretation is consistent

with other paleoenvironmental evidence from northern

Alaska indicating an increase in effective moisture at that

time. For example, the oxygen isotope record from Meli

Lake, located in the Arctic Foothills, shows a shift to

wetter conditions at ~6000 cal yr BP (Anderson et al.,

2001). Similarly, analyses of a sediment core from Lake

Wolverine, located in the Kobuk Valley of northwestern

Alaska, suggest that the lake became deeper at ~5800 cal

yr BP (Mann et al., 2002), The onset of cooler and wetter

conditions after ~5000—4000 cal yr BP is also evidenced

by glacial advances in the Brooks Range and elsewhere in

Alaska (Ellis and Calkin, 1984: Calkin, 1988).

Page 32

Table 1. Mollusk species found in the Red Green Lake sedi-

ment core.

Pisidium (Cyclocalyx) liljeborgi (Clessin, 1886)

Pisidium (Cyclocalyx) nitidum Jenyns, 1832

Pisidium (Cyclocalyx) rotundatum Prime, 1852

Sphaerium (Sphaerium) nitidum Westerlund, 1876

Lymnaea atkaensis Dall, 1854

Valvata sincera helicoidea Dall, 1905

Rising water levels presumably would have affected

the moll community directly by reducing the avail-

ability of shallow-water habitat in the center of the lake.

In addition, the mid-Holocene increase in effective mois-

ture may also have contributed to the decline of mollusks

by triggering other ecological changes. For example, as

the ake became deeper it may ee achieved a depth at

which it could support predatory fish. Lake trout

(Salvelinus namaycush) have been shown to exert strong

controls on mollusk communities in lakes in the Arctic

Foothills (Hershey, 1990; Merrick et al., 1991, 1992;

Hershey et al., 1999), and thus an increase in their popu-

lation as the lake deepened would have exacerbated the

decline in mollusk abundance. The mollusks in Red

Green Lake may also have been affected detrimentally

by changes in water chemistry resulting from climate-

induced ‘changes in aquatic or ‘terrestrial ecosystem pro-

cesses (e.g. , Kling et al., 1992, 2000; Hobbie et al., 1999).

Additional stiches that employ both paleoecological and

paleolimnological approaches are needed to understand

past changes in the linkages between terrestrial and

aquatic ecosystems in the Arctic,

ACKNOWLEDGMENTS

The author thanks Terrence J. Frest, Daniel Gavin,

Linda Brubaker, Feng Sheng Hu, George Kling, Patricia

Anderson, Thomas Brown, Gerald L. Mackie. and the

staff of the Toolik Field Station for their contributions to

this research. The manuscript was greatly improved by

the comments of two anonymous reviewers. This re-

search was funded by National Science Foundation grant

OPP-9615947.

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side, California.

Burch, J. B. 1972. Freshwater Sphaeriacean Clams (Mollusca:

Pelecypc da) of North America. Biota of Freshwater Eco-

systems Identification Manual 3. U.S. Environmental Pro-

tection Agency.

Burch, J. B. 1975. Freshwater Sphaeriacean Clams (Mollusca:

THE NAUTILUS, Vol. 120, No. 1

Pelecypoda) of North America. Malacological Publica-

tions, Hamburg, Michigan.

Burch, J. B. 1989. North American Freshwater Snails. Malaco-

logical Publications, Hamburg, Michigan.

Calkin, P. E. 1988. Holocene glaciation of Alaska (and adjoin-

ing Yukon Territory, Canada). Quaternary Science Re-

wiews 7: 159-184.

Clarke, A. H. 1973. The freshwater mollusks of the Canadian

Interior Basin. Malacologia 13: 1-509.

Clarke, A. H. 1979. Gastropods as indicators of trophic lake

stages. The Nautilus 94: 138-142.

Clarke, A. H. 1981. The Freshwater Molluscs of Canada. Na-

tional Museum of Natural History, National Museums of

Canada.

Delorme, L. D., S. C. Zoltai and L. L. Kalas. 1977. Freshwater

shelled invertebrate indicators of paleoclimate in north-

western Canada during late glacial times. Canadian Jour-

nal of Earth Sciences 14: 2029-2046,

Ellis, J.M. and P. E. Calkin. 1984. Chronology of Holocene

glaciation, central Brooks Range. Geological Society of

America Bulletin 95: 897-912.

Harris, S. A. and E. Pip. 1973. Molluscs as indicators of late-

and post- -glacial climatic history in Alberta. Canadian Jour-

nal of f Zoology 51; 209-215.

Hershey, A. E. 1990. Snail populations in arctic lakes: compe-

tition mediated by predation? Oecologia $2: 26-32.

Hershey, A. E., G. M. Gettel, M. E. McDonald, M. C. Miller,

H. Mooers, W. J. O'Brien, J. Pastor, C. Richards and J. A.

Schuldt. 1999. A geomorphic-trophic model for landscape

control of arctic lake food webs. BioScience 49: S87—897.

Hobbie, J. E., B. J. Peterson, N. Bettez, L. Deegan, W. J.

O’Brien, G. W. Kling and G. W. Kipphut. 1999. Impact of

global change on biogeochemistry and ecosystems of an

arctic fecha: ater system. Polar Research 18: 1-8.

Hu, F.S., L. B. Brubaker and P. M. Anderson. 1996. Boreal

ecosystem development in the Northwestern Alaska

Range since 11,000 yr B.P. Quaternary Research 45; 188—

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Jensen, C., J. G. ]. Kuiper and k. D. Vorren. 2002. First post-

olaciz nl establishment of forest trees: early Holocene veg-

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Kling, G. W., W. J. O'Brien, M. C. Miller and A. E. Hershey.

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Kling, G. W., G. W. Kipphut, M. C. Miller and W. J. O'Brien.

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Merrick, G. W., A. E. Hershey and M. E. McDonald. 1991.

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THE NAUTILUS 120(1):34-36, 2006

Page 34

Mytella charruana (Bivalvia:

in Mosquito Lagoon, Florida

Michelle L. Boudreaux!

Linda J. Walters”

Department of Biology

University of Central Florida

4000 Central Florida Boulevard

Orlando, FL 32816 USA

' [email protected]

* |[email protected]

Mytilidae): a new, invasive bivalve

INTRODUCTION

Biological invasions are recognized as one of the most

serious problems confronting “the integrity of native spe-

cies and ecosystems around the world (Carlton and Gel-

lar, 1993). Invasive species often have no natural enemies

to limit their reproduction and spread; hence, they fre-

quently become established at the expense of the native

species and entire ecosystems (Ellstrand and Schieren-

beck, 2000). In aqui atic ecosystems, extreme examples

include establishment of the zebra mussel Dreissena

polymorpha (Pallas, 1771) in the Great Lakes region of

the United States in the 1980s and the introduction of

the Asian green mussel Perna viridis (Linnaeus, 1758) to

Tampa Bay, Florida in 1998 (G sie et al., L991; Ingrao

et al., 2001). Both of these mussel species: 1) have spread

extensively from the point of introduction via dispersal

larval ae ge s, 2) have cost millions to billions of dollars to

control, 3) have altered biodiversity, primarily by out-

compe ae or overgrowing native species, and +) were so

abundant "ae 1 dissavere d that eradication was not an

option (Ludyanskiy et al., 1999; Fajans and Baker, 2003;

Diggins et al., 2004).

Knowing that early detection and rapid response are

critical to preventing establishment of non-native species

that may cause ecological and economic havoc, we need

to be pro-active when new invasions are reported, This is

especially true when the number of individuals of the

invasive species is persistent, suggesting either multiple

introductions or a resident, breeding population. An ex-

ample of this is presently occurring along the Atlantic

ee of North America in Florida. The bivalve, Mytella

charruana (VOrbigny, 1835), a South American and

Mexican Pacific Coast native, has recently been de-

scribed in low numbers (94 individuals) in the oyster reef

communities of the northern Indian River Lagoon system.

Vytella charruana (VOrbigny, 1835) in Florida

The indigenous range of the mytilid Mytella charruana

extends along the eastern coast of South America from

Venezuela to Argentina (Keen, 1971; Carlton, 1992;

Szefer et al., 1998: Boehs et al., 2004). The charru mussel

also occurs on the eastern Pacific Ocean from Guaymas,

Sonora, Mexico to El Salvador and the Galapagos Islands

(Cardenas and Aranda, 2000),

Mytella charruana resembles the common edible blue

mussel Mytilus edulis (Linnaeus, 1758) in shape. Maxi-

mum recorded shell length is 4.5 em (Szefer et al., 1998).

Its external shell color may vary from light green to black,

and may be uniform or banded in a criss-cross pattern

(Keen, 1971) (Figure 1). Internal shell color is deep

purple (Keen, L971). This tropical mussel is a lagoonal

species that ty pic cally occurs on mudflats, in shallow la-

goons and atti aie to mangrove roots, being able to

withstand salinity variations in the 14-41%o range (Ru-

elas-Inzuna and Paez-Osuna, 2000: Boehs et al., 2004).

Mytella charruana first appeared in the United States

in large numbers in the seawater intake pipe of a power

plant in Jacksonville, FL in 1986 (Lee, 1987; Carlton,

1992). The probable source of invasion was bi a water

from tankers transporting oil from Venezuela (Carlton,

1992). Fortunately, M. charruana never anne estab-

lished, as the founder population was extirpated during

the winter of 1987; this has been attributed to cold tem-

peratures (HI. G. Lee, pers. comm).

Since the 1986-1987 arrival in northern Florida, M

charruana had not been recorded in this location or any

other in the United States until 2004. On August 20,

2004, M. Boudreaux discovered a population of M. char-

ruana in the waters of Mosquito Lagoon, the northern-

most region on the Indian River Lagoon system along the

east coast of central Florida (170 km south of Jackson-

ville), In this environment, M. charruana was initially

found attached to clusters of the eastern oyster Crassos-

trea virginica (Gmelin, 1791) as well as attached to dis-

articulated oyster shells in the intertidal and subtidal

zones within Canaveral National Seashore (28°90.68" W,

80°82.06° N). These waters were then surveyed monthly

for the occurrence of this mussel for the next 13 months.

M. L. Boudreaux and L. J. Walters, 2006

Figure L.

Mytella charruana trom Mosquito Lagoon,

Florida, left (top) and right (bottom) valves, BMSM 34067

length 2.4 cm. Scale bar = 1 em

Systematic surveys were conducted looking at a wide

variety of substrates and places where mussels are known

to thrive, including Hie and bases of the cordgrass

Spartina alterniflora (Loiseleur, 1807), submerged man-

made structures and debris (e.g., dock pilings, PVC pipe,

cinder blocks), natural debris (e.g., driftwood), red man-

grove prop and cable roots, black mangrove pneumato-

phores, clusters of oysters, and disarticulated oyster

shells. For each collected individual, the following infor-

mation was recorded: GPS coordinates, water tempera-

ture (°C), water salinity (%o), mussel length, width and

depth cm), weight (¢) and substrate.

Thirty-nine live individuals of M. charruana were

found between August 2004 and February 2005 and one

dead individual was collected on March 20, 2005. De-

spite continued monthly surveying, no individuals were

found again until August 28, 2005, when 14 live individu-

als were collected. On our last observation date, October

1, 2005, an additional 40 live individuals were collected.

Mytella charruana was found attached to both man-

made debris. especially cinder blocks and PVC, and the

Same small crevice locations On driftwood and oyster

clusters/shells as the native mussel Geukensia demissa

Dillwyn, 1817). In Mosquito Lagoon, water tempera-

ture on dates whe nM. charruana was collected ranged

2 C°. The salinity at collection locations ranged

from 24-36%c on survey dates.

To date, a total of 94 individuals of Mytella charruana

have been found and removed, indicating an invasive

from 15-32

Mean mussel size (cm)

Figure 2... Mean length+standard error (cm) of Mytella char-

ruana collected in Mosquito Lagoon, Florida

population that is still quite small. Although individuals

up to 4.4 cm in length have been recorded in its native

range, the sizes of M. charruana found during one year

in Mosquito Lagoon were consistently small, ranging

from 0.5-2.7 cm (Mean + S.E.: 1.3 + 0.1 em). The mean

length of individuals collected from August 2004—

February 2005 was 1.8 + 0.1 cm, whereas the mean

length for the August-October 2005 collection was 1.1 +

0.1 cm. When compared with one-way analysis of vari-

ance, the mean length of the August 2004-February

2005 individuals was significantly larger (ANOVA: F

32.61; p = 0.0001) (Figure 2). Likewise, the weights of

live individuals collected im August—October 2005 were

significantly lower than weights of live individuals col-

(ANOVA: F = 34.69: p= 0.0003). This

suggests multiple introductions or an established repro-

ductive population, with us locating only newly recruited

individuals. If ecological Conditions: are optim il for con-

tinued survival and establishment of M. charruana, this

population has the potential to greatly crease im num-

bers and out-compete native mussels in the area. Addi-

lected earlier

tional research on this species is warranted to determine

its extent of colonization, source of introduction, envi-

ronmental conditions necessary for establishment, and

impacts on biodiversity. Voucher specimens are cata-

logued at the Florida Museum of Natural History,

Gainesville, Florida (FLMNH 372689, 372691) and The

Bailey-Matthews Shell Museum, Sanibel, Florida

(BMSM _ 34067),

ACKNOWLEDGMENTS

We thank A. ee ie and H. G. Lee for the

identification and J. H. Leal for the photographs of My-

tella charruana. aon of Mosquito Lagoon were un-

dertaken by the spring 2005 Marine Ecology of Florida

class at the University of Central Florida (UCF), P

Sacks, J. Sacks, A. Bz uber, and the C amp Fire USA Sun-

shine Council eee was provided | yy UCF and

Florida Sea Grant. We thank Canaveral National Sea-

een eee nenennnennnneneenen eee

Page 36

THE NAUTILUS, Vol. 120, No. 1

shore for permission to conduct research within Park

boundaries. We thank P. Mikkelsen and an anonymous

reviewer for critically reading and improving the manu-

script.

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Mollusks from Baja California to Peru. 2°¢ Edition. Stan-

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Lee, H. G. 1987. Immigrant mussel settles in Northside gen-

erator. The Shell-O-Gram (Jacksonville Shell Club) 28:

7-9.

Ludyanskiy, M. L., D. McDonald, and D. MacNeill. 1999. Im-

pact of the zebra mussel, a bivalve invader. Bioscience 43:

533-544.

Ruelas-Inzunza, J. R. and F. Paez-Osuna 2000. Comparative

bioavailability of trace metals using three filter-feeder or-

ganism in a subtropical coastal environment. Environmen-

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Szefer, P., J. Geldom, A. Ali, F. Paez-Osuna, A. Ruiz-

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bour, mangrove lagoon of the northwest coast of Mexico.

Environment International 24: 359-374.

THE NAUTILUS 120(1):37, 2006 Page 37

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THE NAUTILUS

Volume 120, Number 2

July 28, 2006

ISSN 0028-1344

A quarterly devoted

to malacology.

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THE@MNAUTILUS

CONTENTS

Volume 120, Number 2

July 28, 2006

ISSN 0028-1344

M. G. Harasewych

George R. Sedberry

Daniel L. Geiger

Roland Houart

Carole M. Hertz

Richard L. Squires

LouElla R. Saul

Richard E. Petit

Rediscovery, range extension, and redescription of Calliostoma torrei

Clench and Aguayo, 1940 (Gastropoda: Vetigastropoda: Calliostomatidae) . . . 39

Sasakiconcha elegantissima new genus and new species (Gastropoda:

Vetigastropoda: Anatomidae?) with disjointly coiled base 2.2... 00 0000.. 45

A review of Typhisopsis Jousseaume, SSO, and Typhisala Jousseaume,

ISS1 (Gastropoda: Muricoidea) of the eastern Pacific 2.2.2.0... 02.000. 52

New buccinoid gastropods from uppermost Cretaceous and Paleocene

strata of California and Baja California, Mexico .. 0... 0000000 ee 66

Authorship of the Ovulidae (Gastropoda) of the Zoology of the Voyage of

thé SAMARANG 002% 4< 2eae3e4e $2 ch beretu bees eee deta toe eas 79

MBLWHOI Library

AUG 05 2006

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THE NAUTILUS 120(2):39—44, 2006

Rediscovery, 1: ange extension, and redescription of Calliostoma

torrei Clench aid Aguayo, 1940 (Gastropoda: Vetigastropoda:

Calliostomatidae)

M. G. Harasewych

Department of Invertebrate Zoology

George R. Sedberry

Marine Resources Research Institute

National Museum of Natural Historv South Carolina Department of Natural

x |

acid : ae Sol Ss

Smithsonian Institution Resource

P.O. Box 37012

Washington, DC 20013-

[email protected]

TOL2 USA

217 Fort Johnson Road

Charleston, SC 29422-2559 USA

[email protected]

ABSTRACT

Two living specimens of Calliostoma torrei were recently col-

lected feeding on bamboo coral (Keratoisis ornata Verrill,

IS7S) east of Jacksonville, Florida, at a depth of $43 m. This

represents a considerable range extension of a species that was

previously known only from the holotype, collected dead off

Matanzas, Cuba, in 704 meters. A re description of the shell and

first descriptions of the operculum, gross anatomy, and radular

morphology are provided; partial sequences of the genes for

16S rDNA and cytochrome ¢ oxidase I have been determined

and added to the GenBank database. These new data suggest

that Calliostoma rugosum Quinn, 1992, from unknown depths

in the Straits of Florida is a synonym of C. torrei. The radula

most closely resembles that of C. yucatecanum Dall, 1881,

suggesting pli cement in Calliostoma sensu stricto according to

the criteria of Clench and Tumer, 1960.

INTRODUCTION

A number of unusual and noteworthy gastropods were

collected during the NOAA Ocean Exploration Cruise

Estuary TO ABYSS, EXPLORING ALONG THE LATITUDE

31-30 TRANSECT, conducted 20 August to 1 September,

2004, off the coasts of South Carolina, Georgia, and

northeastern Florida. Among these were two exception-

ally large specimens of the pleurotomar iid Bayerotrochus

midas (F. M. Bayer

this species northw: are: f from the Little Bahama Bank to

otf the coast of southern Georgia, and two small callios-

tomatid trochoideans. The calliostomatids are members

of a rare and poorly known group of species, character-

ized by a stepped via that are known to inhabit bathyal

depths in the Straits of Florida, throughout the Bz thamas

and the Antilles. southward to Argentina, and eastward to

1965), which extends the range of

the Azores and western Africa. Both specimens appear

closest in morphology to the Calliostoma torrei Clench

and Aguayo, 1940, a taxon known only from the dead-

collected holotype dredged in 704 m off Matanzas, Cuba;

our specimens are smaller and presumed to be imma-

ture. To a lesser extent, they resemble Calliostoma rugo-

sum Quinn, 1992, a taxon that is also based on a single,

dead-collected specimen from the Straits of Florida, but

for which neither depth nor exact location are known.

We provisionally consider all four specimens to be

conspecific and Calliostoma torrei to be the oldest avail-

able name for this taxon. oS this species was pre-

viously known only from dead and damaged specimens,

we provide a detailed de eae of the shell, opercu-

lum, radula, and gross anatomy based on the newly col-

lected specimens. Partial sequences of the mitochondrial

genes for 16S rDNA and cytochrome c oxidase I (CO 1)

have been entered into Genbank, but no se quences for

either of these genes from any species of Calliostoma

were available for comparison at the time of this w riting.

Within the fauna of the western Atlantic, the genus

Calliostoma is unusually diverse, with 94 living species

(Quinn, 1992; Rosenberg, 2005) inhabiting principally

hard-bottom substrates from intertidal to ae bathyal

depths (0—2000 m) at latitudes ranging from 42° N to 55°

(Rosenberg, 2005). Other species inhabit comparable

depths in the Azores and eastern Atlantic. This genus is

thus ideally suited for studies of fine-scale biogeography

and bi thyme tric zonation once sufficient data are accu-

mulated to produce a robust phylogenetic framework.

Institutional abbreviations are: MCZ: Museum of Com-

parative Zoology, Harvard University, Cambridge, Mas-

sachusetts; SERTC/SCDNR: Southeastern Regional

Taxonomic Center/South Carolina Department of Natu-

ral Resources, Charleston, South Carolina; USNM, Na-

tional Museum of Natural History, Smithsonian Institu-

tion, Washington, DC.

Page 40

THE NAUTILUS, Vol. 120, No. 2

SYSTEMATICS (Follows Bouchet et al., 2005)

Clade Vetigastropoda Salvini-Plawen, 1950

Superfamily Trochoidea Rafine sque, 1515

Family Calliostomatidae Thiele, 1924

Genus Calliostoma Swainson, 1840

Calliostoma torrei Clench and Aguayo, 1940

(Figures 1-20)

Calliostoma (Calliostoma) torrei Clench and Aguayo, 1940; 79

80, pl. 14, fig. 5; Clench and Turner, 1960; 59-60, plate ie

Abbott, 1972: 46; Quinn, 1992: 79, figs. SO, SI.

Calliostoma rugosum Quinn, 1992: 102-103, figs. 78-79,

Description: Shell (specimen in Figures 9-12) 22.2

min high, 23.0 mm in maximum diameter, rather thin,

with very narrow umbilicus nearly sealed by parie ti ul fold.

Spire ste pped, weakly concave, spire angle 76.5°. Color

whitish, pale golden yellow between pe riphe »ral keel and

suture, iridescence of aré wonitic nacre visible through

thin calcitic outer layer. Protoconch ( (Figures 14—15)

paucispiral, smooth, glassy, transluscent, increasing from

150 to 500 jum in diameter in about °% whorl (270° ro

tation). Transition to teleoconch marked by flared termi-

nal varix. Teleoconch of 7 1/S dane Step between pe-

ripheral cord and suture concave in first teleoconch

whorl, becoming angular, then anes ssively more con-

vex with increasing whorl number. Base inflated, convex,

evenly rounded. First teleoconch whorl with 21 well-

defined axial ribs and two strong spiral cords (Figure 15,

22 + P3: following cord terminology of Marshall, 1995)

that produce ines sculpture at their i sons

Axial ribs decrease in prominence by 4" teleoconch

whorl, replaced by A NES prominent beads at in-

te me with spiral cords. Ae apical cord (Figures 14,

15, P1) begins as spiral thread in first te eee whorl,

¥+ whorl after the onset of spiral cord P2, expands to

form cord by second teleoconch whorl. Cord P3 remains

at periphery. Fine spiral threads appear between adja-

cent spiral cords at onset of whorl 4, expanding to form

beaded cords by the next whorl. The process repeats,

increasing the number of broad, beaded cords between

P1 and P3 on the body whorl to 6, in addition to 2 fine

and 2 weakly beaded threads. Region between peripheral

cord (P3) ) and suture smoothly concave in teleoconch

ene 1—3, with fine thread appearing at onset of whor!

4 and expanding to form beaded cord by the next whorl,

the process repeating to produce 2 be aded cords and 2

fine threads on penultimate whorl, and 4 beaded cords

and 2 threads on body whorl. A single, smooth cord ap-

pears between suture and PI near end of teleoconch

whorl 7. Base with 21 smooth cords between umbilicus

and periphery. Cords broad near umbilicus, becoming

Figures 1-13.

Calliostoma torrei Clench and Aguayo, 1940. 1-4. Holotype, MCZ 153

W. in 355 fms. [704 m], Harvard-Havana Ixpedition ATLANTIS Station 3485. 1. Apical,

narrower towards periphery. Aperture tangential, with

plane offset from coiling axis by 32°, 11.1 mm wide, 9.6

mm high, outer lip symmetrically elliptical, inner lip with

thickened columella forming an angle of 87° with the

base of previous whorl and 17° with the coiling axis of the

shell. Columella rounded, nacreous inductura nearly

seals very narrow umbilicus.

Operculum: Corneous, thin, transparent, yellowish

amber in color, multispiral, with thin growing edge.

Maximum diameter 8.8 mm.

Gross Anatomy: — Exposed portions of animal yellowish

tan, Foot long, narrow, posteriorly tapering; epipodium

broad, with broad, sc: loped, inhalant and narrow, taper-

ing, enrolled, exhalant neck lobes; 4 epipodial tentacles

per side. Cephalic lappets small, simple, cephalic ten-

tacles long, eye stalks short, with very large, black eyes

Snout large, cylindrical, papillate ventrally with split

ventral lip

Radula (Figures 16-20): Radular ribbon 8.65 mm

long, 1.6 mm wide, with 97 rows of teeth. Rachidian

teeth (Figure 1S, r) simple, triangular, with very finely

serrated edges. Lateral teeth (Figure 18, 1) 7 per side,

with broad attachment area and narrow, simple, curved,

finely serrated along entire outer edges, with or without

fewer, more widely “spaced cusps along distal portion of

inner edge. Innermost marginal teeth ( (Figures 1S, 19,

imm) broad, thick, with 5 short, conical cusps along the

inner distal edge. Marginal teeth become progressively

longe +r and narrower toward the outer e -dge of the radular

hhati: with an increasing number of fine cusps along

the inner, distal por tions of the scythe- shaped feet

Compare inner marginal teeth (Figure 19, im) with outer

marginal teeth (Figure 20, om).

Jaws large, broadly rounded, anterior ends broadly

rounded with short fringe.

Material Examined: Holotype of Calliostoma torrei

Clench and Aguayo, 1940, ATLANTIS Station 3985, off

Matanzas, Matanzas Province, Cuba (23°13' N, 81°22!

W), in 385 fathoms [704 m], MCZ 135165; Holotype of

Calliostoma rugosum Quinn, 1992, JAMEs M. GILLIS

Cruise 7307, Station 13, Straits of Florida, depth and

exact location unknown, USNM 860262; JOHNSON-SEA-

Link IL Dive 3470, Cutthroat Cliff, about 220 km E of

Jacksonville, Florida, (30°17.147' N, 79°20.147" W), in

863 m (SCDNR Collection Number TL0041060), larger

specimen (Figures 9-12) USNM 1069300, smaller speci-

men (Figure 13) SERTC/SCDNR collection $1005.

165, off Matanzas, Cuba, 23°13’ N, 81°22’

2. apertural, and 3. basal views. 4. Detail of

sculpture on last two whorls. 5-8. Calliostoma rugosum Quinn, 1992, holotype re 860262, Straits of Florida, Ames M. GiLis

Cruise 7307, Station 13, depth and exact location unknown. 5. Apical, 6. ape rtural,

. basal views. 8. Detail of sculpture on last two

whorls. 9-12. Calliostoma ene larger of two specimens of collected at Cutthroat c 1th about 135 miles E of Jacksonville, Florida,

30°17.0547" N, 79°20.2514" W, in S70 m, JouNSON-SEA-LINK HT Dive 3470. 9. Apical, 10. apertural, 11. pase! aes, 12

2. Detail of

sculpture on last two whorls. Figure 13. Apertural view of smaller specimen from the same locality. Scale bars = 2 cm for entire

shells, 3 cm for images of sculptural details

M. G. Harasewych and G. R. Sedberry, 2006

Page 42

THE NAUTILUS, Vol. 120, No, 2

Figures 14-15. Calliostoma torrei. 14. Pee and 15. lat-

eral views of protoconch of specimen in Figures 9-12

Habitat and Ecology: The bottom at the Cutthroat

Cliff collection site was sloping at approximately 20°, and

was composed mostly of hard, broken, pavement- like

rock, some carbonate sand and coral ne in mound

formations (up to 10 m high) composed of loosely aggre-

gated coral fragments ead rubble. Scattered dark bios n

manganese-phosphorite pavement and rocks (Popenoe

and Manheim, 2001) were frequently encountered. Bot-

tom temperature was 7.7°C; currents were 0.1—0.2 knots

from the south (180°)

The Calliostoma torrei specimens were living on, and

apparently grazing on, a bamboo coral, Keratoisis ornata.

The coral was attached to a small (20 x 30 em) manga-

nese-phosphorite rock or outcrop that protruded through

light-colored carbonate sand. Stalked crinoids, small

globular and useees sponges, and small erect aS

corals (Lophelia spp.) were common on the dark rocks. /

few cutthroat eels (Synaphobranchus spp.) swam ne ee

The coral branches on which the snails were found (and

some adjacent branches) were devoid of polyps, which

were abund: int and expose «don other branches of the

colony.

DNA Sequences:

drial genes for cytochrome e oxidase I (CO I) (¢

Partial sequences of the mitochon-

yenbank

DQ 314293) and 16S rDNA (Genbank DQ 314294) have

bee n coon ae using standard protocols for DNA ex-

traction, amplification and sequencing.

A search of the Entrez nucleotide database revealed

that only a single sequence for any species of Calliostoma

was present in the database, that for a 302- -bp fragment

of the 28 S rDNA gene of Calliostoma zizyphinum Lin-

naeus, 1758.

Remarks: Calliostoma torrei was described on the ba-

sis of a single, very large (41 mm) but dead-collected

specimen, that has detamorted over time, primarily

through delamination of the outer, calcareous shell layers

from “ihe inner nacreous layers (compare Clench ‘and

Turner, 1960: pl. 40; Quinn, 1992: figs. 80, $1; with Fig-

ures 1—4 herein). Cle nicht and Turner (1960: 60) com-

mented that this species was among the largest in the

western Atlantic and was not closely related to any other

species in the region. They suggested that it was perhaps

closest to the large (to 35 mm), shallow water (30-65 m),

South American species Calliostoma militare Thering,

1907 [as C. amazonicum Finlay, 1930] (Rosenberg,

2005), and, to a lesser extent, to Calliostoma atlantis

Clench and Aguayo, 1940, from comparable depths (603

m) off Mariel, Cuba.

Quinn (1992: 103) described Calliostoma rugosum

from a single, dead-collected and damaged specimen

dredged in the Straits of Florida from unknown depths,

differentiating this from C. torrei based on its smaller

size, broader shell, and differences in the sculpture of the

cords.

The two juvenile, live-collected specimens are inter-

mediate in morphology between the holotypes of C. tor-

rei and C. rugosum. The spire angle of both is closer to

that of C. rugosum, but the sculpture is closer to that of

C. torrei. Distinctions between the four specimens ap-

pear to lie primarily in the relative coarseness of spiral

sculpture, especii ally when compared at corresponding

whorls. We prov isionally consider these specimens to be

conspecific, recognizing that additional material will be

required to better de linesis intraspecific variation over a

range of sizes.

In their review of western Atlantic Calliostoma,

Clench and Turner (1960) did not assign C. torrei to any

subgenus, as neither the morphology of the jaws nor the

radula were known. When compared to ae few species

for which radular morphology has been published, the

radula of C. torre’ appears most similar to that of C.

Laoag i Dall, ISS1 (see Clench and Turner, 1950:

ol. 4, fig. 4), especially in having rachidian teeth without

a broad mae area, narrow lateral teeth, and similarities

in the morphology of the innermost and inner marginal

teeth. Radular and jaw morphology of C. torrei suggest

an affinity with Calliostoma sensu stricto.

Quinn (1992: 103) suggested that both C. rugoswm

and ©. torrei were closely related to the Pliocene C.

caribbeanum Weisbord, 1962, from the Mare Formation

of Venezuela. Based on shell characters, he (Quinn, 1992:

99) hypothesized that the species C. atlantis (603-628 m,

M. G. Harasewych and G. R. Sedberry, 2006

Figures 16-20. Calliostoma torrei. Radula of specimen in Figures 9-12. 16. | Radular ribbon near mid-length. 17. Radular ribbon

anterior to section in Figure 16. Marginal teeth reflected to better reveal rachidian, lateral, and inner marginal teeth. 18. Detail of

rachidian and finely serrated lateral teeth. 19. Lateral view of radular ribbon, showing details of innermost lateral and inner lateral

teeth. 20. Scythe-like outer lateral teeth. im, inner marginal teeth; imm, innermost marginal teeth; 1, lateral teeth: om, outer

marginal teeth; r, rachidian teeth.

off Mariel, Cuba and Great Inagua), C. torrei (including

C. rugosum synonymized herein) (704-870 m, NE

Florida to Cuba), C. dnopherum (Watson, 1879) (640 m,

Recife, Brazil), C. atlantoides Quinn, 1992 (417-589 m,

St. Lucia), C. rota Quinn, 1992 (20-30 m, Brazil), and C.

coronatum Quinn, 1992 (768-805 m, Brazil) form a dis-

tinctive species group. In this group, he also provisionally

included C. cubanum Clench and Aguayo, 1940 (S96 m,

Cardenas, Cuba) and the central and eastern Atlantic

species C. grimaldii Dautzenberg and Fisher, 1896

(1250-2165 m, Azores, Morroco), C. leptophyma

Dautzenberg, 1927 (550-845 m, Azores), C. normani

Dautzenberg, 1927 (599-1600 m, Azores) and C. caroli

Dautzenberg, 1927 (1250 m, Azores). With the exception

of C. rota, all members of this diverse and widespread

group inhabit bathyal depths, most in the 500—L000 m

range, with depths increasing eastward.

ACKNOWLEDGMENTS

Funding for submersible operations was provided by a

grant (No. NAOROAR4600055, G.R. Sedberry, Principal

Investigator) from the National Oceanic and Atmo-

spheric Administration Office of Ocean Exploration. Su-

san DeVictor, of the SERTC/SCDNR laboratory, iden-

tified the coral on which the Calliostoma were collected.

The SERTC staff assisted in processing the NOAA-OF

samples and submersible pilot Tim Askew, Jr. (Harbor

Branch Oceanographic Institution) assisted in collec-

tions. This is Contribution Number 582 of the South

Carolina Marine Resources Center, and Smithsonian

Marine Station at Fort Pierce Contribution Number 648.

LITERATURE CITED

Abbott, R. T. 1974. American Seashells, 2"! Edition. Van Nos-

trand Reinhold Co., New York, 663 pp., 24 pls.

Bouchet, P., J. Fryda, B. Hausdorf, W. F. Ponder, A. Valdés

and A. Warén. 2005. Working Classification of the Gas-

tropoda. Malacologia 47: 239-397,

Clench, W. J. and Aguayo. 1940. Notes and descriptions of new

deep-water Mollusca obtained by the Harvard-Habana

Expedition off the coast of Cuba. HI. Memorias de la

Sociedad Cubana de Historia Natural 14(1): 77-94

Clench, W. J. and R. D. Turner. 1960. The Genus Calliostoma

in the Western Atlantic. Johnsonia 4(40); 1—SO.

Dautzenberg, P. 1927. Mollusques provenant des campagnes

scientifiques du Prince Albert I de Monaco dans Océan

Atlantique et dans le Golfe de Gascogne. Résultats des

Page 44

THE NAUTILUS, Vol. 120, No. 2

Campagnes Scientifiques accomplies sur son yacht par le

Prince Albert I, Prince de Monaco 72: 1-400.

Dautzenberg, P. and H. Fischer. 1896. Campagnes scienti-

fiques de S. A. le Prince Albert I" de Monaco. Dragages

effectués par /Hirondelle et par la Pricesse Alice, 1SSS—

1895. I. Mollusques gasteropodes. Memoires de la Société

Zoologique de France 9: 395-498.

Hickman, C.S. and J. H. McLean. 1990, Systematic Revision

and Suprageneric Classification of Trochacean Gastro-

pods. Science Series of the Natural History Museum of

Los Angeles County 35: vi + 1-169 pp.

Marshall, B. A. 1995. A Revision of the Recent Calliostoma

Species of New Zealand (Mollusca: Gastropoda: Tro-

choidea). The Nautilus 108: 83-127.

Popenoe, P. and F.'T. Manheim, 2001. Origin and history of

the Charleston Bump-geological formations, currents, bot-

tom conditions, and their relationships to wrecktfish habi-

tats on the Blake Plateau. In: Sedberry, G. R. (ed.) Island

in the stream: oceanography and fisheries of the Charles-

ton Bump. American Fisheries Society Symposium 25, Be-

thesda, pp. 43-93.

Quinn, J. F.. Jr. 1992. New species of Calliostoma Swainson,

1840 (Gastropoda: Trochidae) and notes on some poorly

known species from the western Atlantic. The Nautilus

106: 77-114.

Rosenberg, G, 2005. Malacolog 4.0.1: A database of Western

Atlantic marine Mollusca. [WWW database (version

4.0.1)] URL http://data.acnatsci.org/wasp |

THE NAUTILUS 120(2):45-51, 2006

Sasakiconcha elegantissima new genus and new species

(Gastropoda: Vetigastropoda: Anatomidae ?) with disjointly

coiled base

Daniel L. Geiger

Santa Barbara Museum of Natural History

Invertebrate Zoology

9559 Puesta del Sol Road

Santa Barbara, CA 93105 USA

[email protected]

ABSTRACT

Sasakiconcha elegantissima, a new genus and species from the

Western Pacific is described. The specimens are most similar to

members of Anatomidae (Anatoma, Thieleella) with a periph-

eral selenizone, but close the slit to a foramen, and lack a

columella because of a disjointly coiled base resulting in an

umbilical trough. Additionally, an aberrant specimen of Scis-

surella morretesi Montouchet, 1972, which shows repeated clo-

sure of the slit to form foramina, is discussed with respect to the

evolutionary pathway from the ancestral open slit to its modi-

fication as a single or a series of foramina.

Additional Keywords: Protoconch, teratology, biodiversity,

scanning electron micrographs, SEM

INTRODUCTION

The study of the scissurellid grade has revealed a signifi-

cant number of new taxa over the last few years, from

species to family-level (e.g., Marshall, 1993; 2002; Kase

and Kano, 2002: Bandel, 1998; Geiger, 2003; 2006; Gei-

ger and Jansen, 2004a, b). With these new taxa, more and

more discrete shell types have been discovered. Shells

with flat spires (Satondella Bandel, 1998; Maxellella Ban-

del, 1998), the calyptraeiform Depressizona Geiger,

2003, and even the new slit-less forms such as Troglo-

concha Kase and Kano, 2002, expanded on the more

common trochiform shell morphologies.

Until recently, the above forms had been classified in

Scissurellidae: however, recent molecular studies have

shown that Scissurellidae is a grade (Geiger and Thacker,

2005). More specifically, Scissurellidae (Scissurella, Si-

nezona, Satondella, Incisura) and Anatomidae (Anatoma,

Thieleella) do not form a clade, whereas other lineages

Sutilizoninae, Temnocinclinae, Depressizoniné 1c ) have

yet to be included but are likely not to be part of either

of the sec juenced lineages (see G eiger and Thacker, 2005

for discussion).

The careful study of existing museum collections has

been the main driving force behind these discoveries.

Here, I report on some specimens found in the collection

of the Muséum National d’Histoire Naturelle, Paris that

exhibit yet another new shell morphology. Additionally, a

note on an abnormal specimen of Scissurella morretesi

Montouchet, 1972, shows that the transition from single

slit to multiple foramina can occur even in modern spe-

cies.

MATERIALS AND METHODS

Specimens were mounted on double-sided carbon tabs

(Ted Pella, Redding, California), sputter-coated with

gold i ina Cre ssington 10S auto sputter coater with rot: ary-

planetary stage, and viewed in a Zeiss EVO4O XVP

(Zeiss, Cambridge, Uk) scanning electron microscope.

Specimens of the new species were cleaned with a wet

artist brush and a tungsten needle; the s specimens ap-

peared to be too fragile to withstand ultrasonic cleaning.

The number of specimens is indicated by the number

separated by a comma from the museum acronym (see

below). Protoconch whorls are counted following Geiger

(2003).

Institutional abbreviations used in the text are: AMS:

Australian Museum Sydney, New South Wales, Austra-

lia. BMNH: The Natural History Museum, London,

Great Britain; LACM: Natural History Museum of Los

Angeles County, Los Angeles, California, USA; MNHN

Muséum National dts Naturelle, Paris, France:

NMNZ: Museum of New Zealand Te Papa Tongarewa,

Wellington, New Zealand; UMUT: University Museum,

University of Tokyo, Japan; USNM: United States Na-

tional Museum, Smithsonian Institution, Washington,

DC, USA.

SYSTEMATICS

Vetigastropoda Salvini-Plawen, 1980

? Anatomidae McLean, 1989.

Sasakiconcha new genus

Page 46

THE NAUTILUS, Vol. 120, No. 2

Type species: Sasakiconcha elegantissima new spe-

cies.

Differential Diagnosis: Anatoma and Thieleella

share the position of the slit at the periphery, but the slit

remains open, and the suture is always below the lower

keel of the selenizone, even in the most depressed spe-

cies, A. paucispiralia (Bandel, 1998).

Description: Shell depressed biconical, whorls rapidly

increasing in size, no nacre. Protoconch sometimes par-

tially covered by suture of subsequent whorl. Whorls not

touching at base forming umbilical trough; no columella.

Selenmore and foramen at periphery. Suture mostly at or

above upper keel of selenizone of previous w horl. Um-

bilicus wide. Animal unknown.

Etymology: Sasaki-: honoring Takenori Sasaki of the

University Museum of the Univ. ersity of Tokyo for his

significant contributions to the systematics of basal gas-

tropods. -concha: Latin for shell. Sasaki’s shell. Cendex

feminine.

Remarks: —Sasakiconcha is tentatively placed in Anato-

midae because of the peripheral position of the seleni-

zone. That position was shown to discriminate between

major scissurellid/anatomid lineages in an exploratory

phylogenetic analysis of shell chances (Geiger, 2003).

The most likely alternative hypothesis is that "Sasdhicon:

cha represents yet another independent lineage of slit-

bearing, small vetigastropods.

Many species of Anatoma show conv erging margins of

the slit, but in no species do they fuse and enclose a

foramen. Sasakiconcha is not based on senile specimens

of Anatoma, as the foramen is found over a wide range of

shell sizes (4-8 mm). Not a single specimen of Anatoma

or Thieleella with a closed foramen is known.

Sasakiconcha must form its foramen by a different

process compared to most Sinezona or Sukashitrochus.

Sasakiconcha closes the foramen from fairly early growth

onward, i.e., at about half-maximum size, in contrast to

most scissurellids, which close the foramen at about 4/5

of maximum size. It implies that, with growth, the ante-

rior margin of the foramen is resorbed, while shell ma-

terial is added at the apertural margin, leaving a foramen

of approximately constant proportion over a wide range

of shell sizes (4-S mm). In contrast, juveniles of Sinezona

and Sukashitrochus have an open slit, which only closes

once the animal is fully grown. Only Sinezona globosa

Geiger, 2006, known from the two type specimens and

USNM 500687 (Caribbean locality data most likely

wrong), also closes the foramen in not fully grown speci-

mens (Geiger, 2006). Comparison to other foramen-

bearing small vetigastropod genera of the scissurellid

grade (Sutilizona, Temnocinclis, Temnozaga, Depressi-

zona) is not possible due to the scarcity of material.

The overgrowth of the selenizone by the shoulder of

the subsequent whorl is unique among Recent Vetigas-

tropoda, but is known from some fossil genera ( Brookes-

Knight et al., 1960): e.g., Agniesella Cossmann, 1909

Portlockiellidae), Umbotropis Perner, 1903 (Gosseletin-

idae), and Catachisma Branson, 1909 (Gosseletinidae).

The overall shell morphology is most similar to the cae-

nogastropod genus Cochliolepis Stimpson, 1858 (Vit-

rinellidae), though this latter misses the selenizone in-

dicative of a vetigastropod placement for Sasakiconcha.

Sowerby (1820-1834) illustrated Scissurella elatior Sow-

erby, 1831, on an unnumbered plate with the selenizone

overgrown by the subsequent whorl. This character is not

discussed as the species was introduced as a nomen et

figura, without indication of where it was found. Sowerby

(1842, 1852) placed it as a Recent species occurring off

the coast of Britain, from where only Anatoma crispata

(Fleming, 1828) is known. It is thus most likely that Sow-

erby’s (1820-1834) figure was stylized and is erroneous

with respect to the overgrowth ‘of the selenizone. The

whereabouts of the type eienal of Sowerby’s species is

unknown.

Sasakiconcha elegantissima new species

(Figures ]—2)

Differential Diagnosis: The cited generic characters

will readily identify S. elegantissima. The most similar

species is Anatoma paucispiralia (Bandel, 1998) de-

scribed from Satonda, Indonesia. Shared characters in-

clude the overall biconical depressed shell shape, the

protoconch lacking an apertural varix, and the short te-

leoconch I. The preeerreh is half the size in A. pau-

cispiralia (150 wm vs. 325 jum), teleoconch I is longer in

A. paucispiralia (0.125 whorls vs. 0.05), the suture is

below the lower keel of the selenizone of the previous

whorl (i.e., the selenizone is not obscured), the sculpture

of the shoulder has distinct axial and spiral components,

the base has reticulate sculpture, and the slit is open

although the aperture of the illustrated oe (Bandel,

1998: pl. 14, figs. 2-4) is slightly chipped.

Description: Shell biconical-depressed, Sinum-

shaped, to $.5 mm (holotype); thin, fragile. Protoconch

285-325 jum (mean = 311 wm, n = 4, 325 pm in holo-

type), 0.75 whorls, flocculant sculpture (mostly eroded in

holotype), no apertural varix, apertural margin convex;

sometimes partially covered by suture of * subsequent

whorl. Teleoconch I <0.05 whorls. Teleoconch IT up to

1.75 rapidly expanding whorls. Shoulder slightly convex,

early teleoconch II with axial cordlets only, interstices

irregular with tiny pits; after 0.5 whorls, axial cordlets

with approximi ately one dozen nodules, nodules becom-

ing spirally elongated with strength of axial cordlets de-

creasing; after 1.25 whorls, sculpture transformed into

approximately 50 spirally arranged, irregularly spaced,

nodular cordlets; interstices with finest axial growth

marks. Base with fine, stepped, longitudinally undulat-

ing, spirals, reaching just into uppermost portion of um-

bilo! cavity, approximately 30 in 4 mm specimen, ap-

proximate ly 40 in S mm specimen, Umbilicus wide, walls

steep, smooth; umbilical wall not connecting with previ-

ous whorl, forming spiral trough between umbilical wall

and previous whorl produced be inner surface of shoul-

der. Selenizone at periphery, usually hidden by suture of

D. L. Geiger, 2006

Figure 1.

scale bar prot sconch = 200 Jz.

subsequent whorl: slit closed anteriorly to form foramen,

foramen formed in specimens =4 mm. In fully mature

specimens (holotype), last 0.125 whorls descending. Ap-

erture oblong, D-shaped, roof overhanging. Animal un-

known.

Type Material: Holotype (MNHWN type collection);

paratypes: MNHN, 1 (Figure 2: left), 477-493 m,

BORDAU1, DW1432, off Fiji, 17.333° S, 178.733° W;

MNHN, 1 (Figure 2: middle), 300-302 m, BORDAU2,

DW 1523, Eua, Tonga, 21.300° S, 175,000° W; MNHN,

2 (one illustrated in Figure 2: right), 391-421 m,

BORDAU2, DW1537, haut-fond, 21.683° S, 175.317:

W: UMUT RM29213, 1; NMNZ M.273290, 1, 700-707 m,

BORDAU], DW1455, Fiji, 19.050°S, 17S.500°W.

Type Locality: 970-953 m, BORDAU2, DW1531,

Eua, Tonga, 21.200°S, 174.933°W.

Holotype of Sasakiconcha clegantissima new genus and new species, MNHN type collection. Scale bar shell = 2 mm;

Etymology: Elegantissima: elative of Latin adjective:

extremely elegant referring to the overall aspect of the

shell.

Other Material Examined: — Fiji: MNHN, 1, 700-707 m,

BORDAU1, DW1485, 19.050° S, 178.500° W: Tonga:

MNHN, 1, 456-510 m, BORDAU2, DW1509, Ton-

gatapu, 21.083° S, 175.367° W; MNHN, 2, 578 m,

BORDAU2, DW15S5, Vava'u group, 18.550° S$, 173.950!

W: French Polynesia: MNHN, 1, 600-620 m,

BENTHAUS, DWISS9. east of Rap, 27.613° S, 144.262° W.

Distribution: Western Pacific (17-

W), 800-951 m [all shells only |.

°S, 144-175

Remarks: Sasakiconcha is not a morphological end-

point in a series of undescribed Anatoma species The

MNHN collection contains 23 undescribed Anatoma and

Page 48 THE NAUTILUS, Vol. 120, No. 2

Figure 2. Paratypes of Sasakiconcha elegantissima new genus and new species. First row: apertural view: second row: lateral view

showing closure of slit to foramen; third row: umbilical view; fourth row: apical view; fifth row: protoconchs. Left: MNEIN, 1, 477-493

m, off Fiji, 17.333° S, 178.733° W middle: MNHN, 1, 300-302 m. Eua, 21.300° S, 175.000° W: right: MNHN, 2, 391-421 m,

haut-fond, 21.683°S, 175.317° W. Only in one of the smaller specimens on the left the slit is still open. In the other two, which are

approximately half the size of the holotype, the slit is already closed to a foramen, Scale bars shells = 1 mmy scale bars protoconchs

100 xm: TL: extent of teleoconch I

D. L. Geiger, 2006

Page 49

Figure 3. Juvenile Haliotis rubra Leach, 1S14, from a hatch-

ery. A short slit-less teleoconch I follows the protoconch. Te-

leoconch II immediately forms a series of foramina. Scale bar =

1 mm.

Thicleella species from the Indo-Pacific awaiting detailed

study through electron microscopy. None of these even

remotely approaches Sasakiconcha.

The umbilical trough is a character not known from

any other scissurellid sensu lato or anatomid and is to my

know ledge a novel configuration in gastropods; its func-

tion is unclear. Umbilical modifications have been shown

to function as brood pouches in Larocheinae (Marshall,

1993).

Sasakiconcha elegantissima is a fairly large anatomid

spe cies, and one with 1a diameter of S mm dees not even

qualify as a micromollusk, which usually is defined as

shells smaller than 5 mm. Despite its size, it is not the

largest known anatomid. A specimen of Anatoma eu-

glypta ( (Pelseneer, 1903) from Antarctica is 10.5 mm in

diameter (Zelaya and Geiger, unpubl. data).

The protoconch of Sasakiconcha elegantissima is as

largest known of any scissurellid/< aaremid species by ¢

factor of approximately two. It implies that rather ae

eggs are formed and that the number of eggs is relativ ely

small, as has been observed in other scissurellids and

anatomids [Sinezona rimuloides (Carpenter, 1865);

Anatoma euglypta: Geiger, unpubl. data]. This is quite

unlike the more familiar gonad condition in the large,

broadcast spawning Ui iain oda with millions of eggs

(e.g., Bevelander, 1988). Large eggs themselves do, how-

ever, not entail brooding or parental care. Eggs may also

be deposited onto any available hard substrate as shown

for Scissurella jucunda Smith, 1890 by Strasoldo (1991)

REPEATED CLOSURE OF THE SELENIZONE

The selenizone with associated slit or foramen/foramina

is typically found in Vetigastropoda, including Scissurel-

lidae sensu lato (not in Larocheinae), Anatomidae, Pleu-

rotomariidae, Fissurellidae, and Haliotidae. whereas in

other families it is absent (Trochoidea, Lepetodrilidae,

Peltospiridae, Neomphalidae). A sinus is formed in Seg-

uenziidae, which had prompted Locard (1898) to include

Seguenzia in Scissurellidae, whereas this latter genus is

placed today in its own group besides Vetigastropoda.

That the open slit is the ancestral condition can be

argued both based on data from the fossil record as well

as from the ontogeny of Recent species. The open slit is

found in the various pleurotomarioid families since the

Paleozoic (Brookes-Knight et al., 1960; Harasewych,

2002), as well as in juveniles of Sinezona and Sukashitro-

chus, which eventually closes the slit to form a foramen

at maturity (Geiger, 2003). Mature shells can be recog-

nized by the distinct descending last quarter whorl of the

adult shell. Haliotidae have modified the shell in a par-

ticular fashion in that they repeatedly open and close the

slit to form a series of foramina (Delhaes, 1909). Most

likely, the closure of the foramen lends more stability to

the apertural margin of the shell. Similar shells of Scis-

surella with an open slit break more easily than those of

Sinezona with a closed foramen. As Haliotidae usually

live in high energy regions of the intertidal and shallow

subtidal ( (Shephe rd, 1973), a strengthening of the aper-

tural margin throughout their life makes the repeated

formation of foramina advantageous. The underlying

mantle has a single, anteriorly open, slit, covering all

open foramina (Crofts, 1929), while in foramen be -aring

Fissurellidae all stages from open mantle slit to fused

mantle skirt can be observed (McLean and Geiger,

1998S).

Intermediate forms are not widely known. Delhaes

(1909) discussed the keyhole-shaped double foramen of

the fossil pleurotomarioideans in the genera Ditremaria

and Phragmostoma. Ontogenetically, Haliotidae imme-

diately form a series of holes after a short slit-less teleo-

conch I (Bevelander, 1988: fig. 4.1; Sasaki, 199%: fig. 28;

fig. 3), unlike in Sinezona, where the juvenile has an

open slit, which is closed only once at maturity (Geiger,

2003).

A single specimen of an aberrant Scissurella morretesi

is shown here (Figure 4). Two specimens of the species

were available in eo with the original descrip-

tion of Montouchet (1972). As it is virtually eae to

legally obtain loans from Brazil at this time, no further

material was available. No further exemplars of the spe-

cies were found out of a total of 29,566 specimens ex-

amined (major collections include: AMS, BMNH,

LACM, MNHN, NMNZ, USNM: many smaller collec-

tions not listed). Despite other abnormalities observed

such as senile closure of foramen in Scissurella spp., the

condition of the $. morretesi specimen discussed here is

unique. The first SEM illustrations of the species are

proy ided here.

As for all Scissurella species, usually the slit remains

open throughout life. In the particular specimen, the _

was closed and re-opened three times successively.

does not appear to have been caused by damage to [

shell; the shoulder and base of the shell in the vicinity of

the closure show no growth irregularities. It rather seems

Page 50

THE NAUTILUS, Vol. 120, No. 2

Figure 4

Scissurella morretesi Montouchet, 1972. Left: four views of normal specimen. Scale bar shell = 1 mm; scale bar

protoconch = 100 zm, Right: apical view of aberrant specimen with evidence of three-fold successive closure of slit to foramen (bold

arrows). Square shows portion of shell shown enlarged below; inset: Enlarged view of second closure; connected arrows show onset

and end of closure; MNHN, 295 m, SE, Brazil, 18.983° S, 37.833° W. Scale bar shell = 1 mm; scale bar inset = 500 wm.

to have been caused by some sort of mutation/develop-

mental abnormality. The specimen provides an illustra-

tion of a possible pathway from the ancestral open slit, to

irregularly repeated closure of the slit, to patterned clo-

sure of the slit as seen in Haliotidae.

ACKNOWLEDGMENTS

This study was made possible by a Visiting Curatorship

from MNHN, and NSF grant MRI 0402726 to Henry

Chaney, Michael Caterino, and Daniel L. Geiger. I

would like to thank all staff at MNHN for their hospi-

tality and assistance during the ste ly. Jame s McLean and

Diego Zel: wva offered constructive criticism that helped

to improve the manuscript. Richard Petit provide d de-

tails on the dating of some of Sowerby’s works.

LITERATURE CITED

Bandel, K. 1998. Scissurellidae als Modell fiir die Variations-

breite einer natiirlichen Einheit der Schlitzband-

schnecken (Mollusca, Archaeogastropoda). Mitteilungen

des Geologisch-Palaontologischen Instituts der Universitit

Hamburg $1: 1-120

Bevelander, G. 1988. Abalone Gross and Fine Structure. The

3oxwood Press, Pacific Grove, California. SO pp

Crofts, D. R. 1929. Haliotis. Liverpool Marine Biology Com-

mittee Memoirs 29: 1-174, pls. 1-S

Delhaes, W. 1909. Beitriige zur Morphologie und Phylogenie

von Haliotis Linné. Zeitschrift fiir induktive Abstam-

mungs- und Vererbungslehre 2: 1-55, pls. 2-3

Geiger, D. L. 2003. Phylogenetic assessment of characters pro-

posed for the generic classification of Recent Scissurel-

lidae (Gastropoda: Vetigastropoda) with a description of

one new genus and six new species from Easter Island and

Australia. Molluscan Research 23: 21-83.

Geiger, D. L. 2006. Eight new species of Scissurellidae and

Anatomidae (Mollusca: Gastropoda: Vetigastropoda) from

around the world, with discussion of two new senior syn-

onyms. Zootaxa 1128: 1-33.

Geiger, D. L. and P. Jansen. 2004a. Revision of the Australian

species of Anatomidae (Gastropoda: Vetigastropoda).

Zootaxa 435: 1-35.

Geiger, D. L. and P. Jansen. 2004b, New species of Australian

Scissurellidae (Mollusca: Gastropoda: Vetigastropoda)

with remarks on Australian and Indo-Malayan species.

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THE NAUTILUS 120(2):52-65, 2006

Page 52

A review of Typhisopsis Jousseaume, 1880, and Typhisala

Jousseaume, 1851 (Gastropoda: Muricoidea) of the

eastern Pacific

Roland Houart!

Institut royal des Sciences naturelles

de Belgique

Rue Vautier, 29

1000 Bruxelles, BELGIUM

Carole M. Hertz”

Santa Barbara Museum of Natural History

2559 Puesta del Sol Road

Santa Barbara, CA 93105 USA

ABSTRACT

Typhisopsis Jousseaume, 1880, and Typhisala Jousseaume,

ISSI, are redescribed and the esa species reviewed. Ty-

phisopsis coronatus (Broderip, 1833) is considered as the senior

synonym of T. quadratus (Hinds, 1843), Murex siphoniferus

Lesson, 1844, and Typhis martyria Dall, 1902. In addition,

Typhisopsis carolskoglundae new species is described from

western Panama. Its range extends to Sonora, Mexico. The new

species is compared with Typhisopsis coronatus, Typhisala

grandis (A. Adams, 1855) and T. clarki (Keen and Campbell,

1964).

The four species have different shell morphology, especially in

the number, height, and breadth of the spiral cords, the mor-

phology of the axial ribs and the siphonal canal. The proto-

conchs of the four species are illustrated for the first time.

Lectotypes are designated for Murex siphoniferus and for Ty-

phisala grandis.

INTRODUCTION

A few months ago, the senior author received a lot of

specimens of “Typhis” species from Costa Rica for iden-

tification. At first sight that lot was separated into two

species: Typhisopsis coronatus and Typhisala grandis.

However, after more careful examination and discussion,

we began a complete re-evaluation of the whole group

occurring in the eastern Pacific. Typhisopsis coronatus

was described from a sub-adult specimen. Another spe-

cies, previously identified as a dwarf form of Typhisala

grandis by D’Attilio (1987) and D’Attilio and Hertz

(1988S) was re-examined. It is here described as a new

species of Typhisopsis.

Keen (1944), Gertman (1969), Radwin and D’Attilio

(1976), Vokes (1988), and other authors classified all the

Typhis-like species in the Muricidae, subfamily Typhinae

Research Associate

- Associate

Cossmann, 1903. D’Attilio and Hertz (1988) included

the species related to Typhis in the family Typhidae [as

in Sacco (1904) and Garrard (1963)] and split it into two

subfamilies: Typhinae and Tripterotyphinae D’Attilio

and Hertz, 1988. Their study was based on morphologi-

cal differences of shell and radular structure. The validity

of Tripterotyphinae is not questioned herein. Subse-

quent authors, however, chose to retain Tripterotyphi-

nae, as well as Typhinae, as a subfamily of Muricidae

instead of Typhidae (Vokes, 1989a, 1989b, 1996: Houart.

L991, 1994, 3009: Absalao and Santos, 2003). DNA work

currently under way elsewhere (Oliverio and Houart, in

a ) for Typhinellus labiatus (Cristofori and Jan, 1832

= Typhis sowerbyi H. and A. Adams, 1858) as well as

oe DNA work may help clarify the taxonomy in this

group. Radular terminology follows Houart (1991) based

on D’Attilio and Hertz (1988).

Abbreviations used in this study are: BM(NH): Natu-

ral History Museum, London, Great Britain; CAS: Cali-

fornia Academy of Sciences, San Francisco, California,

USA; IRSNB: Institut royal des Sciences naturelles de

Belgique, Bruxelles, Belgium; LACM: Natural History

Museum of Los Angeles County, Calif fornia, USA;

MNHN: Muséum national d’Histoire naturelle, Paris,

France ; MZCR: Museo de Zoologia, San José, Costa

Rica; SBMNH- Santa Barbara Museum of Natural His-

tory, C alifornia, USA; SDNHM: San Diego Natural His-

tory Museum, California, USA; SUPTC: Stanford Uni-

versity Pale eee Type Collection, Stanford, USA;

USNM: ee al Muse sum of Natural History, W ‘ashing-

ton, D.C. USA; CJH: Carole and Jules Hertz ( ‘ollection,

San Diego, California, USA; CS: Carol Skoglund Collec-

tion, Phoe nix, Arizona, USA; RH: Roland Houart Col-

lection, Landen, Belgium; RZ: Ricky Zandali ¢ Io

Abaco, Bahamas: spec.: live-taken specimen; dd spec.

empty shells, dead collected.

SYSTEMATICS

Subfamily Typhinae Cossmann, 1903

Description: Shells with aperture entire, usually with

projecting peristome and no sulcus: usually four or more

R. Houart and C. “Her ‘rtz, 2006

varices per whorl, either wing-like or with axial swellings:

rounded, ventrally sealed anal tubes originating from

adapical spiral cord, only current anal tube functional,

older tubes sealed: siphon: i canal ventrally sealed with

left side ove rlap] ying right side.

Genus Typhisopsis Jousseaume, 1SSO

Type Species: = Typhis coronatus Broderip, 1833, east-

ern Pacific, by original designation.

Description:

adult with varical flange, frilled abaperturally. Varices

heavy, rounded, rope -like. Shoulder spines broad-based,

triangular, flat, tapering inwards to a sharp point. Parti-

tion connecting apertural shoulder spine and last teleo-

conch whorl. Anal tubes situated near preceding varix.

Siphonal canal bri vad. ventrally sealed. Left side of canal

broadly overlapping right side

Typhisopsis coronatus (Broderip, 1833)

Figures 1-15, 16, 44-46, 56-57, 61-62, 67)

Typhis coronatus Broderip, 1833: 178; Sowerby, 1841, pl. 200,

figs 3.4

Typhis quadratus Hinds, 1843: 18; Hinds, 1844: 10, pl.3, figs 3, 4.

Murex siphoniferus Lesson, 1844: 168.

Typhis martyria Dall, 1902; 550; Dall, 1908, pl.15, fig.11.

Typhis (Typhisopsis) coronatus.—kKeen, 1944: fig. 17; Keen

1971: 540, fig. 1051 right [type figure of T. quadratus|

Abbott, 1974: 192 (2017

Typhisopsis coronatus.— D Attilio, 1975: 57, text fig.; D’Attilio,

1976: 2S, text figs: Radwin and D Attilio, 1976: 212, pl. 32

figs. 10-12: Abbott and Dance, 1982: 157, text fig

Shell with four varices; apertural varix of

D’Attilio, 1987: 34-35, figs 7-11; D’Attilio and Hertz

LOSS: 21, fig. LO9

Typhis coronatus.—Kaicher, 1978: card 1595

Typhis (Typhisopsis) grandis —keen, 1971, 540, fig. 1052 [not

Typhisala grandis (A, Adams, 1855)|

Not Typhis (Typhisopsis) coronatus Keen, 1971: 540, fig

L051 (left) not Broderip, 1833 [= Typhisala grandis (A

Adams, 1855) |

Description: Shell up to 40 mm in length. Biconic,

heavy; spire high, acute; protoconch off-white, of 2.75

rounded whorls, first protoconch whorl pustulose re-

maining whorls smooth; terminal lip thin, raised, curved

(Figures 44-46). First four teleoconch whorls smooth,

elossy, with no microse ulpture; intritacalx cove ring pen-

ultimate and final whorl with microsculpture consisting

of regular gouge-like pits (Figure 57). Shell with six

weakly shouldered. nodose teleoconch whorls. Last

whorl broad; suture moderately impressed and mostly

obscured. Axial sculpture of four broad, rounded, rope-

like varices, each nee a broad, flat, triangular,

strongly inward-bent, shoulder spine at intersection with

P2. On last whorl apertural varix broadly expanded, edge

reflected dorsally, frilled abapertur: ally with posterior

edge extending into a sharp

shoulde ‘r; anal siphon ee) ) adjace nt to prece ding varix

and appressed : partition connecting the varix of previ-

ous whorl. Final anal siphon long and dorsally directed.

Spiral sculpture on three

raised spiral eve (P3-P6) becoming less prominent at

, dorsally incurved spine at

earliest varices of heavy

the anterior end, intervarical spiral sculpture faint or ab-

sent. Siphonal canal long, broad (Table 1), ventrally

Figure 1.

Morphology of the spiral cords in Typhisopsis coronatus Broderip 1833

Page 54

THE NAUTILUS, Vol. 120, No. 2

Table 1. Terminology used to describe the spiral cords (based

on Merle 1999, 2001)

P Primary cord

S Secondary cord

Pl Shoulder cord

P2—P6 Primary cords on the convex part of the

teleoconch whorl

sl-s6 Secondary cords on the convex part ot the

teleoconch whorl

Example: sl = secondary cord between PI and P2;

s2 = secondary cord between P2 and P3, ete.

ADP. adapertural primary cord on the siphonal canal

MP median primary cord on the siphonal canal

ABP abapertural primary cord on the siphonal canal

ads adapertural secondary cord on the siphonal canal

ms median secondary cord on the siphonal canal

abs abapertural secondary cord on the siphonal canal

sealed along its length with 5 cords (probably ADP, ads,

MP, ABP, abs), tapering with long, backwards facing,

sealed tube at distal end. Sutural line strongly shifted to

the right. Aperture white, moderately large, Smeets sub-

Gireulae. entire, forming an erect peristome appresse od

posteriorly. Intritacalx chalky, off-white, deeply and ir-

regularly pitted. Shell color white to cream with brown

suffusions on varices, tips of shoulder spines, and tip of

siphonal canal; two to five brown spots on edge of aper-

tural lip, sometimes none, on occasional specimens these

spots also on leading side of varices. Operculum corneus,

light brown with 13-15 concentric lamellae and terminal

nucleus. Radula (Figure 67) with rachidian tooth bearing

a long central cusp, and on each side 3 or 4 slightly

shorter, narrower lateral denticles of varied strength. and

length or erratically fused, and a broad lateral cusp, slightly

shorter than central cusp. Lateral teeth sickle shape id,

broad.

Type Material: = Typhis coronatus: Ecuador, Salango,

holotype BM(NH) 1966447; Typhis quadratus: Costa

Rica, Golfo de Nicoya, and Ecuador, Bahfa de Guay-

aquil, dredged from a muddy bottom 7 to 1S fms (13-33

m) (Golfo de Nicoya is here de ssignated as type locality),

holotype BM( NH) 1844.6.7.35; Murex siphoniferus:

Mexico, Acay yulco, Guerrero, | lectotype MNHN (here

selected); Ty) his ah a Golfo de California, off San

Pedro Martir, 26 m, sand , holotype U ad 130629.

Other Material Examined: Mexico, Bahia Magca-

lena, Baja California Sur, 1 spec. (SBMNH); San Felipe,

Baja California, | spec. (SBMNH); Bahfa de los Angeles,

Baja California, dredged 27 m, | spec. (SDNHM 23216

figured in Radwin ani D Attilio, 1976]|); off Isla San

Marcos, Baja California Sur, divers, 12 m on Spondylus

princeps, | spec. (CS); Isla Carmen, Baja California, 1 dd

spec. (SDNHM); Baja California Sur,

dredged 35-40 m, 3 spee. (CS); dredged 30-45 m, 3

spec. Cy :

SDNHM)

Isla Danzante,

Jahia ( foncepcion, Baja California, | spec.

Cabo Tepoca, Sonora, dredged 20-300 m,

sand bottom with rocks, 5 spec. (CS); Guaymas, Sonora,

dredged, 24 m, Ispec., (RH); 1 spec. dredged ( SDNHM

43812);55 m 1 dd spec. (MNHN); Bahia Bacochibampo,

Sonora, dredged 42 m, 1 spec. (MNHN); dredged 18-27

m, 2 spec. (CJH); 150 spec. (SBMNH): Sonora 1 spec.

(MNEHIN), 2 spec. (RH); off_ Bahia San Carlos, Sonora,

19-38 m, pe (IRSNB IG 26.158); dredged 18-22 m,

98 spec. (CJH); dredged 15-30 m , 14 spec. (CS); 4 spec.

31 m (SDNHM 90837): drec dged 100 m, 3 spec.

(SDNHM 71920): off Tetas de Cabras, dredged 60-100

m, 7 spec. (SDNHM 92766); Tenacatita, Jalisco, 2 spec.

(SBMNH): Manzanillo, Colima, near harbor entrance,

15-22 m, 1 spec. (RH); dredged 10-30 m, 3 spec. (CS);

dredged 31 m, 4 spec (SDNHM); dredged 100 m,

3 spec. (SDNHM); dredged 60-100 m, 7 spec.

(SDNHM): 16 spec. (SBMNH): Salina Cruz, Oaxaca, 5

spec. (SBMNH); Costa aie Playas del Coco, Guana-

caste dredged 24 m, 4 spec. ); dredged 24-36 m, 4

spec. (CJH); dredged ao m, 2 spec. (SDNHM

90774): 1 dd spec. (SDNHM 91506); 1 spec. (SBMNH):

16 spec. (SBMNH); Panama, 21 spec. (SBMNH); Isla

Santa Catalina, dredged 5-8 m, 5 spec. (RE); Boca de la

Honda, 7°27'N, 80°51'W, in white sand, 2 spec. (RH);

SE Isla Rancheria, Golfo de Chiriqui, dredged 15-30 m,

1 spec. (SDNHM 90775); Isla Cébaco, off Golfo de

Montijo, Veraguas, in silty sand and rubble, 6-9 m, 1

spec, (MNHN); Bahia Montijo, Veraguas, 19 m, 1 spec.

(RH); Arenas de Quebro, dredged 19-38 m, 1 spec.

(RH); Isla Venado, 1 dd spec. (SDNHM 64236); Ecua-

dor, Islas Gal&pagos, Isla San Salvador, close to Bahia

James, 30-35 m, coll. D.R. Shasky, 1 spec. (SBMNH

366002)

Distribution: San Felipe, Baja California and Cabo

Tepoca, Sonora, Mexico, to Guayaquil, Ecuador and Islas

Galapagos, 542 m.

Remarks: Subadult specimens of T. coronatus (Fig-

ures 11-12, 62), as in the type, have an incompletely

formed apertural lip which gives the shell a narrower and

more slender outline. The first four teleoconch whorls

(to a length of approximately 15 mm) of T. coronatus are

smooth and shiny without the microsculpture evident on

mature shells. In juvenile specimens the siphonal canal is

also narrower, having amore centrally situated sutural line.

Of the two syntypes of Murex siphoniferus (MNHN),

one is Typhisala grandis, the other is conspecific with

Typhisopsis coronatus. As first revisers, and in order to

maintain the name T. grandis as it was illustrated by

recent authors, we designate that latter specimen of

Murex siphoniferus as lectotype ( (Figures 6-7). Two of

the three syntypes of Typhisala grandis ( (A. Adams, 1855)

are specimens of Typhisopsis coronatus (see under that

species for further explanation). The holotype of T.

quadratus (BM(NH)) has the siphonal canal broken , but

the overall shape of the shell is quite the same as

intact specimen illustrated by Hinds (1S44: pl. 3, figs

3-4). No other specimen was located in the type imine

rial,

R. Houart and C. M. Hertz, 2006

Figures 2-15. Typhisopsis coronatus (Broderip, 1833), 2-3. Ecuador, Salango, holotype BM(NH) 1966447, 26 mm (photo P

Crab. BMNH). 445. Typhis quadratus Hinds, 1843, Costa Rica, Golfo de Nicoya (herein designated as type locality), holotype

BM(NH) 1544.6.7.35, 17.5 mm (photo P. Crab, BMNH). 6-7. Murex siphoniferus Lesson, 1544, Mexico Acapulco, Guerrero

lectotype MNHN (here designated), 27.9 mm (photo D. Brabant, MNHN). 8-9. Typhis martyria Dall, 1902 holotype USNM

130629. Golfo de California, off San Pedro Martir, 27.6 mm. 10. Typhisopsis coronatus Tetas de Cabra, Guaymas, Sonora, Mc

19.4 mm, CJH photo P. Sadeghian protoconch illustrated Figures 43-45), 11-12. Panama, Santa Catalina, 5—S m, RH

13-15. West Panama. Boca de la Honda, 7°27’ N, 80°51’ W, in white sand, RH, 23.56 mm

Page 56

THE NAUTILUS, Vol. 120, No. 2

Typhisopsis carolskoglundae new species

(Figures 17-25, 47-49, 59, 63)

Typhisala grandis. —D Attilio, 1987: 32, figs. 1-6; D’Attilio and

Hertz, 1988: 72-73, figs. 10S a-e [not Typhisala grandis

(A. Adams, 1855)].

Description: Shell up to 30.9 mm in length, slender,

heavy. Spire high, 40-48% of total shell length. Proto-

conch off- shite. conical with 2.7 rounded protoconch

whorls, first whorl pustulose, remaining whorls smooth,

ending with delicate, thin, weakly erect and curved ter-

minal lip; (Figures 47-49): ): teleoconch with up to 5 or 6

broad, strongly shouldered whorls. Suture impressed,

partly Ghseuned: axial sculpture of first three teleoconch

whorls with sharp varices; remaining teleoconch whorls

consisting of four strong, broad, rounded varices with

remains of sharp lamina visible on varices of last whorl:

each varix with a broad, flat, long, strongly inwards

pointed triangular, flat spine at adapical extremity touch-

ing previous whorl, Anal tube adjacent to leading edge of

varix and appressed to previous partition. Varices

broader and swollen at shoulder periphery. Apertural

varix with a moderately broad, recurved, sinuous flange,

a slightly broad, long, somewhat curved spine at posterior

edge, spine atti ache d to previous whorl by an axially

erooved partition densely sculptured spirally; lip edge

rounelly curving into siphonal canal approximately 1/4

from the end.

Apertural varix with strong spiral cords beginning at its

outer lip and over-ridden by strong axial fimbriation;

outer recurved edge of abapertur: al dde squamose, Spiral

sculpture of raise J. heavy, primary cords on varices. Last

whorl with 15 cords (P1, P2, s2, P3, s3, P4, s4, P5, s5, P6,

followed by rounded cords on siphonal canal, probably

ADP, ads, MP, ms and ABP), forming a broad expansion

at adapical extremity of apertural varix. Pl with sealed,

rounded anal tube near preceding varix. P2 with flat,

triangular spine at intersection with varices. Intervarical

sculpture faint or absent under magnification (20x).

Sealed rounded anal tube appressed to partition on pre-

ceding varix, forming an angle of approximately SO-90°

with axis of shell. Spiral Gone ending as short, strongly

backward curved spines at edge of apertural varix.

Aperture subcircular, forming a continuous peristome,

erect at outer lip and anterior portion of columellar lip

and appressed ee sriorly. Siphonal canal comparatively

short, broad, ventrally sealed. Sutural line strongly

shifted to the right, le ft side of canal broadly overlapping

right side. Intact specimens ending with canal dorsally

reflected and curved to the right at extremity.

Fresh, not overly cleaned specimens, with a simple,

chalky off-white intritacalx showing occasional axial

striae. Protoconch off-white with white pustules on first

whorl. Teleoconch color white to cream with brown on

shoulder spines and anal tubes; 2-5 brown spots on edge

ol outer apertural lip anc often visib le On leading edge of

pre vious varices. One brown spol midw: ay intervie arically

on each varix except the apertural varix, as in Typhisala

grandis. Operculum light brown with 12-14 concentric

lamellae and terminal nucleus. Radula not examined.

Type Material: Holotype: SDNHM 90773, Costa

Rica, Playas del Coco, Guanacaste, on mud bottom, 24—

37 m; Paratypes: Costa Rica, le del Coco, Guana-

caste, on mud bottom, 24-37 m, 1 paratype (SDNHM

93558); on mud ey, 24-37 m, 1 paratype (CJH);

dredged 12-25 m, 9. (CS); 2 spec., 18-27 m in coarse

sand, broken shell ecole SBMNH 3504: 31); 1 paratype

(SBMNH 359436): 5 spec., 9-30 m (SBMNH 359435): 1

paratype (USNM 1084298); 1 paratype (LACM eagle

Isla Negritos, Golfo de Nicoya, 3 paratypes, 21-27 m,

coarse seach broken shell (SBMNH 359430): Panama,

Arenas de Quebro, Veraguas, 19-37 m, 1 paratype (RH);

Boca de la Honda,7°27' N, 80°51’ W, in white sand, 1

paratype (BM(NH) 20050371); 1 paratype ogee UCR

6153); 1 paratype (MNHN Moll 6991); 2 paratypes

(RH); off Isla Gobernadora, intertidal, 1 paratype (RH);

off Isla Venado, Bahia Panama in muddy sand at low tide,

| paratype (IRSNB IG 26.817/566): Bahfa Chiriqui, 74

m, 1 paratype (IRSNB IG 28.466/567); Contadora, Per-

las Archipelago, ee de Panama, 4 paratypes, 15-30 m

(SBMNH 359432): 1 paratype (SBMNH 359429); Ecua-

dor, Isla oe ie Islas Galapagos, 1 paratype (SBMNH

359433).

Other Material Examined: Mexico, Bahfa San Car-

los, Sonora, 31 m, 1 spec. (RH).

Type Locality: Costa Rica, Guanacaste, Playas del

Coco, on mud bottom, 24—37 m.

Distribution: Sonora, Mexico to Isla Venado Bahia de

Panama, Panama, from the intertidal to 74 m and Islas

Galapagos, Ecuador.

Etymology: This species is named in honor of Carol

Skoglund of Phoenix, Arizona, author and specialist in

Panamic mollusks, who first collected the type material

of this species and donated the holotype and a paratype

to the SDNHM. She has also donated additional

paratypes of this species from her private collection to

the USNM and to the LACM.

Remarks: = Typ/iisopsis carolskoglundae differs from T.

coronatus in having a broader siphonal canal with an

expanded varical flange, extending almost to the tip of

the siphonal canal; flange sinuous midw: ay to the anterior

end, roundly curving into the siphonal canal, approxi-

mate ‘ly 1/4 from the end and constricted at the base. It

has a relatively smooth shell surface under a chalky,

simple intritacalx, compared to the squamose, pitted in-

tritacalx in T. coronatus. The axial ribs are usually

broader at the shoulder, and there are more numerous

and slightly narrower spiral cords on the last teleoconch

whorl, 15 in contrast to 11-12 in T. coronatus. The va-

rices appear broader in coronatus and the spiral cords are

less pronounced in the new species.

The holotype of Typhisopsis carolskoglundae was il-

R. Houart and C. M. Hertz, 2006

Mexico, Colima, Manzanillo

Species of Typhisopsis and Typhisala. 16. Typhisopsis coronatus (Broderip, 1833

22 m, coll. RH, 24.38 mm. 17-25. Typhisopsis carolskoglundae new species 17-18. Costa Rica, Playa del

Figures 16-29.

near harbor entrance, 15-22

Coco, Guanacaste. on mud bottom, 24-37 m holotype SDNHM 90773, 17.8 mm photo K. Barwick). 19. Paratype SDNHM

93555$17.1 mm photo K. Barwick), 20-22. Western Panama, Boca de la Honda, 7°27’ N, 80°51’ W, in white sand, paratype RH

30.1 mm. 23. Western Panama, Boca de la Honda, 7°27’ N, S0°51' W, in white sand, paratype MNHN Moll 6991, 18.9 mm. 24.

i. Bahia San Carlos, 31 m, RH, 26 mm. 25. Paratype SBMNH 359436, type locality, 16.6 mm (photo P. Sadeghian

26-27. Bahia Panama, Isla Venad

22 mm photo P Crab, BMNH

Mexico, Sonor

llustrated Figures 46-48). 26-29. Typhisala clarki (Keen and Campbell, 1964

prot ncl

protoconch illus

holotype CASIZ 064667 [ex-SUPTC 9724], 22.5 mm photo G Metz). 28-29. Paratype BM(NH

SESS RD

Page 58

THE NAUTILUS, Vol. 120, No. 2

lustrated in D’Attilio (1987: figs. 1 and 2 [left]) and

D’Attilio and Hertz (1988: fig. 108, ae), as a dwarf Ty-

phisala grandis, but Typhisopsis carolskoglundae differs

from Typhisala grandis in having a smaller, narrower

shell at maturity and fewer and Bronder spiral cords on

the last teleoconch whorl (15 in Typhisopsis carolsko-

glundae in contrast to 20-23 on Typhisala grandis). Ty-

phisopsis carolskoglundae also has broader, shorter, flat

anal shoulder spines, typical of Typhisopsis and nar-

rower less sharp axial ribs with the flange on the aper-

tural varix more strongly constricted at he base of the

siphonal canal than in Typhisala ¢ erandis.

Typhisopsis carolskoglundae differs from Typhisala

clarki in having broad, rounded varices, rather than the

sharp varices in T. clarki, in having anal tubes adjacent to

the leading edge of varices rather than midway between

varices, and in having brown spots on the aperture edge

and on previous varices, which are lacking in Typhisala

clarki. The new species is similar to grandis and clarki in

having an expanded, strongly fimbriate varical fl: ange and

a senoctlh shell under a simple chalky intritacalx.

Genus Typhisala Jousseaume, 18S]

Type species: = Typhis grandis A. Adams, 1855, eastern

Pacific, by original designation.

Description: Shell with four varices; apertural varix of

adult with broad apertural varical flange extending to

almost the tip of siphonal canal, fimbriate abaperturally;

other varices sharp, narrow. Shoulder spines narrow;

spines of three abapical teleoconch whorls narrow,

twisted, curved inwards; partition connecting apertural

spine and last teleoconch whorl. Anal tubes situated near

preceding varix, pressed against preceding partition. Si-

phonal canal broad, ventrally sealed; left side of canal

overlapping right side.

Typhisala ce Keen _ Campbell, 1964)

(Figures 26-29, 30-34, 50-52, 58, 64, 68)

Typhis (Typhisopsis) clarki Keen and Campbell, 1964: 48, pl. 9,

figs. 15, 19, 23

Typhis (Typhisopsis) clark. —Keen, 1971: 540, fig. 1050; Ab-

bott, 1974: 192 (2016).

Typhis clarki.—Gemmell, 1974: 100-103, 5 figs.; Kaicher,

1980: card 2508

Typhisala clarki—D Attilio, 1975: 58, text fig.; Radwin and

D Attilio, 1976: 211, pl. 31, fig. 5; Gemmell and D’Attilio,

1979: SS—93, figs. 1-8: D’Attilio and Hertz, 1988: 20, fig.

LOT.

Description: Shell up to 27 mm in length, biconic,

lamellate, delicate. Spire moderate [ hk 39-42% of

total shell length; protoconch of 2 5 off-white, smooth,

convex, conical whorls, first whorl pustulose, remaining

whorls smooth (Figures 50-52); teleoconch with six an-

gulate, strongly shouldered whorls. Suture impressed,

partly obscured by succeeding whorl, Axial sculpture of

teleoconch consisting of four high, sharp, narrow, lamel-

late varices, each swith a slightly open, long, twisted, in-

ward curved spine adapically, each varix “composed of

two appressed laminae, one on the receding edge ex-

tending further than the one on leading edge, thie two

forming the slightly open a at shoulder Anal siphon

see forming an angle of approximately 85—90° with

axis of shell, placed approximately midway between va-

rices and appressed to previous partition, last tube only

complete, long, open. Apertural varix flange- like,

strongly fimbriate, dorsally reflected as shallow, open

spinelets at outer edge. Ape rtural spine broad, triangular,

weakly or strongly Foveally curved, connected to last te-

leoconch whorl by a broad, high, lamellate, partition. All

other spiral cords ending as short, weakly or strongly

backward curved, open spinelets at edge of apertural

varix. Spiral sculpture faint, where present, 12-14 inter-

varical lines visible on some specimens under magnifica-

tion (LOx) corresponding to open spinelets at outer ap-

ertural edge. On fresh specimens, not over-cleaned, in-

tritacalx sculpture of simple axial striae present. P1 with

sealed, tapering, anal tube. P2 ornamented with long,

upwards curved spine at intersection with varix.

Aperture small, ovate, forming a continuous peris-

ie erect at outer lip and anterior portion of columel-

lar lip. Siphonal canal long, broad, ventrally sealed, ta-

pering to a slender, dors: lly curved tube at distal end.

Sutural line weakly shifte d to right, left side of canal

weakly overlapping right side. Color of protoconch off-

white with white pustules on first whorl; teleoconch

brown, cream or yellowish-white (sometimes with all

three colors on one shell), having darker brown anal

tubes, shoulder area, shoulder spines, and tip of siphonal

canal. Apertural flange often white even on brown shells.

On fresh specimens, not over-cleaned, intritacalx sculp-

ture of simple axial striae present. Operculum corneus,

light brown with 12 or 13 concentric lamellae and ter-

minal nucleus, Radula (Figure 68) with rachidian tooth

bearing a long central cusp, and on each side 2 shorter,

narrower, lateral denticles of approximately same

strength and length, and a broad lateral cusp, slightly

shorter than central cusp. Lateral teeth sickle- shape od,

broacl.

Type Material: Holotype: Panama, Isla Venado, Bahia

Panama, Panama CASIZ 064667 (formerly SUPTC

9724): 1 paratype CASIZ (formerly SUPTC 9725); |

paratype BM(NH) 1964433.

Other Material Examined: Mexico: Pta. San Felipe,

Baja California, intertidally, in mud and rocks, 28 spec.

(CJ); intertidal, in mud between rocks, 7 spec. (CS):

intertidal, —1.8 m tide, 19 spec. (SDNHM = SI10SS);

spec., intertidal, -1.5 m tide (SDNHM 53452, figured

in Radwin and D’Attilio, 1976); intertidal, 9 spec.

(SDNHM $2868); 31 spec., intertidal (SBMNH); Bahia

la Cholla, Sonora, intertidal in sand over rocks, 7 spec.

(CS); intertidal, —1.5 m tide, 2 spec. (SDNHM 90835):

R. Houart and C. M. Hertz, 2006 Page

Figures 30-43. Species of Typhisala. 30-34. Typhisala clarki (Keen and Campbell, 1964). 30-32. Panama, off Isla Cébaco

dredged in 6-9 m, silty sand and rubble, RH, 22.8 mm. 33. Bahia Panama, Isla Venado, under stones at low tide, RH, 22.3 mm. 34.

Mexico, San Felipe. Baja California, (CS), 25.0 mm (photo P. Sadeghian). 35-43. Typhisala grandis (A. Adams, 1S55), 35-36,

California, lectotype BM(NH) 1974470 (here selected), 37.9 mm (photo P. Crab, BMNH). 37-39. Mexico, Sonora, San Carlos

Punta Doble, 20-22 m, in rubble, RH, 33.7 mm. 40. T. grandis, SBMNH, Mexico, Sonora, Guaymas, Bahia San Carlos, 15 m, 18.0

mm photo P. Sadeghian). 41-43. Costa Rica. Province of Puntarena, [sla Ballena, Canton de Osa, offshore, between 12-25 m, RH

24.0 mm

Page 60 THE NAUTILUS, Vol. 120, No. 2

Figures 44-55. Protoconchs. 44-46. Typhisopsis coronatus (Broderip, 1833), 47-49. Typhisopsis carolskoglundae new species

50-52. Typhisala clarki (Keen and Campbell, 1964). 53-55. Typhisala grandis (A. Adams, 1855). Scale bars: figures 44, 47, 50, 53

0.5 mm; other figures = 200 zm. All photos by D. Geiger.

5 spec. (SBMNIT); Panama: off Isla Cébaco, dredged IG 26

17); intertidal, 9 spec. (CS): Isla Gobernadora,

in 6-9 m, silty sand and rubble, 1 spec. (RH); Isla Ve- intertidal, 1 spec. (SDNHM 93342); Bahfa de Panama

nado, under stones at low tide, 1 spec. (RH); 21 spec. intertidal (SDNHM 62897): Isla Pedro Gonzales, Islas

(SBMNH); in muddy sand at low tide, 1 spec. (IRSNB Perlas, 2 spec. (SBMNH).

R. Houart and C. M. Hertz, 2006

Figures 56-60. Detail of spines and intritacalx. 56-57. Typhisopsis coronatus (Broderip, 1833). 58. Typhisala clarki (Keen anc

Campbell, 1964). 59. Typhisopsis carolskoglundae new species 60. Typhisala grandis (A. Adams, 1855). All RH. Scale bars = 2 mm

Distribution: San Felipe, Baja California and Bahia la

Cholla, Sonora, Mexico to Panama, intertidally to 6 m.

Remarks: The elegant lamellae of the varices, the

smooth or nearly smooth shell surface, and the lack of

bre WL spots on the apertural lip and previous varices of

T. clarki distinguish it from the other three species

Typhisala grandis (A. Adams, 1855)

Figures 35-43, 53-55, 60, 65-66, 69)

Murex siphoniferus Lesson, 1544: 168 (paralectotype MNHN)

see under Typhisopsis coronatus).

Typhis grandis A. Adams, 1855: 42, pl. 27, fig. 4.

Typhis Typhisopsis crandis.—Keen, 1944: 54, fig. 18; Abbott,

1974: 192 (2018).

Typhisala grandis.—D Attilio, 1975: 57, 1 text fig.; Radwin and

D Attilio, 1976: 211, pl 29. fig. 1: Abbott and Dance, 1982

15S. 1 text fig

Typhis Typhisopsis coronatus.—Keen, 1971: fig. 1051 (left

not T. coronatus Broderip, 1833

Not Typhis (Typhisopsis) grandis —Keen, 1971; 540, fig. 1052

= Typhisopsis coronatus

Not Typhisala erandis—D Attilio, 1987: figs. 1-6; D’Attilio

and Hertz, 1988: fig. 108, a-e (= Typhisopsis carolsko-

clundae new species

Description: Shell up to 37.9 mm in length, broad,

heavy. Spire low or moderately high, angulate; proto-

conch off-white of 2.5 rounded, smooth whorls, first

whorl not pustulose (Figures 53-55); teleoconch of up to

S1X strongly shouldered whorls, broadly rounded on last

two whorls. Suture deeply impressed, obscured by suc-

ceeding whorls. Axial sculpture from first teleoconch

whorl to antepenultimate whorl consisting of four low,

sharp, varices, becoming broadly rounded at periphery of

penultimate and last whorl with a sharp flange extending

out from the varix becoming fimbriate on its leading

edge; each varix with a moderately slender, curving spine

at its adapical extremity. Immediately adjacent to the

spine is a deep indentation anteriorly, next to which the

anal siphon (tube) rests against the former partition. Fi-

nal anal siphon (functioning tube) long, at approximately

90° to the shell axis. Apertural varix broadly expanded,

constricted at anterior end; outer edge, reflected and

densely fimbriate; connected to previous teleoconch

whorl by a broad, high, densely lamellose partition. Ap-

ertural spine short, sharply dorsally curved

Spiral sculpture of sharp, raised primary cords, sec-

ondary cords visible under magnification (10x). Last

whorl, including siphonal canal with 22-23 cords. P1

with sealed, rounded anal tube, P2 ornamented with

short spine at intersection with varix. Other cords cannot

be quantified. Spiral cords of approximately same mag-

nitude, more obvious on varices, ending as short, strongly

backwards curved, short spine at outer edge of apertural

varix.

Aperture round-ovate, forming a continuous erect

peristome; outer lip with 3-5 brown spots often appear-

mg as well on leading edge of varices, brown spot also

appearing on each intervarical area; Siphonal canal long

f ~iAD

Page 62

THE NAUTILUS, Vol. 120, No. 2

Figures 61-66.

broad (Table 1) ventrally sealed. Sutural line strongly

shifted to the right, left side of canal broadly overlapping

right side. Microsculpture of thin, low, curved, lamellae

covering the canal and weakly overlapping varices, except

apertural varix. Intritacalx off-white, sometimes thick,

with occasional axial striae

Color milky-white to lavender, light tan, tan or almost

entirely dark brown with brown colored spines, anal

Siphonal canal. 61-62. Typhisopsis coronatus (Broderip, 183:

carolskoglundae new species 64. Typhisala clarks ( (Keen and Campbell, 1964). 65-66. Typhisala grandis (A. Adams, 1855). 65. Adult

specimen. 66. Juvenile. All RH. Scale bars = 0.5 mm.

). 61. Adult specimen. 62. Juvenile. 63. Typhisopsis

tubes, and shoulder area. Other brown spots between

spiral cords, on outer lip of aperture, on leading edge of

varices, not intervarically and occasionally on sutural line

of siphonal canal. Operculum corneus, light brown, with

11-12 concentric lamellae and terminal nucleus.

Radula (Figure 69) with rachidian tooth bearing a long

central cusp, and on each side 3-4 short, narrow, lateral

denticles of varied strength and length, occasionally fused

R. Houart and C. M. Hertz, 2006

Figures 67-69. Radulae (from D’Attilio and Hertz, 1955S).

67. Typhisopsis coronatus (Broderip, 1833), Islas Galapagos,

Isla Jervis. 68. Typhisala clarki (Keen and Campbell, 1964),

Mexico, Baja California , San Felipe. 69. Typhisala grandis (A.

Adams, 1855), Mexico, Bahia de Banderas. Unknown magnifi-

cations.

or split, and a long, broad lateral cusp slightly shorter than

central cusp. Lateral teeth sickle-shaped, broad.

Type Material: BM(NH) 197470, 3 syntypes here se-

lected as lectotype and paralectotypes (Figures 35-36),

“California” (see Remarks).

Other Material Examined: Mexico: Punta Doble,

San Carlos, Sonora, 20-22 m, in rubble, Ispec. (RH); 2

dd spec. 20-21 m diving. (SDNHM 90834); 15-21 m, 4

spec. (CS); divers, 20-21 m in rubble, 2 dd spec.

(SDNHM): off Bahia San Carlos, Sonora, dredged 31-75

m, 12 spec. (CS), 3 spec. (C JH); dredged 60 m, 2 spec.

(CJH); dredged 3-5 m, 4 spec. (SDNHM 76475); 64

spec. (SBMNH): 3 mi SE San Antonio, Guaymas, So-

nora, dredged 100 m, 2 dd spec. (SDNHM 80764): Man-

zanillo, Colima, 20 spec. (SBMNH): >in 6 m, (SDNHM

23215 [as quadratus|); Costa Rica: Isla Ballena, Puntar-

enas, offshore, 12-25 m, 4 spec. (RH); 4 spec., ( RZ); Isla

Ballena, Canton de Osa, Puntarenas, offshore, between

12 and 25 m, 10 spec. (subadult, dd) (RH); 3 spec.

C]JH): 2 spec. (RZ); Panama: 10 spec. (SBMNH); Isla

Gobernadora 1 spec. (MNHN); extreme low tide, in

muddy sand near rocks, 1 spec. (RH); low tide, in muddy

sand, 1 spec. (RH): Isla Cébaco, dredged in 37 m, sé ind

and gravel bottom, 1 spec. (RH); 62-93 m, 1 spec.

(IRSNB IG 26.817); 37-52 m, 1 spec. IRSNB IG

27.037); Isla Santa Catalina, dredged 5-8 m, 1 spec.

(RE); Isla Venado, intertidal, 1 spec. (CS). Ecuador: Islas

Galapa wos, Isla San Salvador, close to Bahia James, 30-

35 m, coll. D.R. She isky, 1 spec. (SBMNEL 366003).

Distribution: Punta Doble,

Panama and the Islas

62 m.

Sonora, Mexico to

Galépagos, Ecuador, intertidally to

Remarks: Of the three syntypes of Typhis grandis

housed in BM(NH), two are Typhisopsis coronatus, the

other is Typhisala orandis. In order to maintain the sta-

tus of T. grandis as it was illustrated by recent authors

(except Keen, 1971), one syntype specime n is here des-

ignated as the lectotype (Figures 35-36). The type local-

ity is doubtful because, to our knowledge, no specimen

has ever been re ported from California since then. Many

times in old publications “California” is also used for

“lower California” now known as Baja California which is

part of Mexico. It is probably the case here. Subadult

forms of T. grandis (Figures 40-43, 66) differ from the

adult form, in having a narrower, lower partition, a

rounded apertural varix without varical flange, a nar-

rower, more acute, siphonal canal, with the ventral su-

tural line being more central. In adults the left side of the

canal extensively overlaps the right side (Figure 65).

Discussion: Both Typhisopsis and Typhisala seem to

be closely related, and it seems questionable whether

both are separate or not. However, several differences

have been detected so far: In Typhisopsis the varical

spine is flat, broadly triangular and strongly inward bent

(Figures 56, 59), compared to the longer, weakly twisted

spine in Typhisala (Figures 57, 60) and the apertural

varix of Typhisala is fimbriate and broadly expanded

whereas the apertural varix of Typhisopsis is narrower

and more rounded with heavy raised spiral cords. The

question is whether or not such differences are signifi-

cant enough to warrant separate genera. Mi iybe DNA

work weld be useful in this ] sapeoulae case.

ACKNOWLEDGMENTS

We are most grateful to M. G. (Jerry) Harasewych and

Paul Greenhall (USNM) for the oan of Typhis martyria

Dall, 1902; to Kathie Way and Phil Crabb (BMNH) for

digital images; to Virginie Héros and Delphine Brabant

(MNHN) for help in searching literature and specimens

from the collection of MNHN and for digital images; to

Paisley Cato and Laura Halverson (SDNHM) for loan of

the two type specimens of Typhisopsis carolskoglundae;

to George Metz , Novato, California, for digital images of

the holotype of Talityphis clarki at CAS; to Kelvin Bar-

wick (City of San Diego EMTS Lab) for digital images of

the holotype and paratype of Typhisopsis carolskoglun-

dae; to Daniel Geiger (SBMNH) for SEM photographs

of the protoconchs of the four species; to Patricia Sade-

ghian (SBMNH) for digital images of the complete speci-

oF a &

Page 64

THE NAUTILUS, Vol. 120, No. 2

Table 2. Characters of Eastern Pacific Species of Typhisopsis and Typhisala.

Character

Character states

Typhisophis coronatus

Typhisopsis

carolskoglundae

Typhisala grandis Typhisala clarki

1. Protoconch

Last teleoconch

whorl®

1. Shoulder spine

2. Anal tube

position

3. Number of spiral

cords (siphonal

canal included)

4. Axial ribs

5. Apertural varical

flange

Siphonal canal®

1. Mean:

breadth/length

2. Position of the

sutural line

Intritacalx

2.75 white, conical,

rounded whorls,

pustulose on first whor!

Moderately broad

Flat, broad, not twisted at

base, sharp, incurved at

acute end

Tubes adjacent to leading

edge of preceding varix

and appressed to

previous partition,

10-11, broad

Broad, rope-like rounded

varices, apertural varix

with varical flange

Edge reflected; weakly

squamose, spiral cords

obvious, more strongly

squamose on outer

portion

Moderately broad, long,

tapering

6.08/7.76 mm

Broadly ove rlapping the

right side

Dinstinctive, chalky,

off-white, deeply

irregularly pitted

2.66 rounded whorl

pustulose on first

Broad

Flat, not twisted, long,

broad at base, cu

at acute end

Tubes adjacent to leading

edge of varix and

appressed to previous

partition

13-15, narrow, broadly

spaced

First three teleocon

whorls with sharp

g with small, thin,

varices; remaining

whorls broad, rounded;

apertural varix Wi

broad varical flange,

Ss, 2.5 cream-colored,

2.75 white, smooth, conical

rounded whorls, pustulose

on first whorl

Broad

rounded whorls,

none pustulose

whorl

Broad

Weakly twisted, broad base,

long, open at distal end

Weakly twisted, long,

broad at base, bent

dorsally

rved

Tubes adjacent to

preceding varix,

appressed to

preceding partition

Tubes approximately midway

between varices, appressed

to preceding partition

21-22, narrow, broadly

spaced on siphonal

canal

12-14, only visible on sharp

edge of axial ridges,

otherwise smooth

ch Broad, rounded:

abapical portion

Angulate, sharp, formed by

two appressed lamellae;

apertural varix expanded.

erect lamellae;

th apertural varix with

broad varical flange

roundly curving into

siphonal canal

approximately 0.2

before distal e saa

Broad, recurved;

fimbriate on its o

pe yrtion, roundly

curving into sphe

canal

Moderately broad

$.05/7.S5 mim

Broadly overlapping right

side

Chalky, off-white, si

with occasional axial

striae

Broadly expanded, Broad, reflected as shallow

uter recurved: strongly open spinelets; strongly

fimbriate on its fimbriate on its outer

vical outer portion, portion

strongly constricted

at anterior end

Broad Papering, long

9.41/9.57 mm 4.91/7.62 mm

Broadly overlapping

Strongly overlapping the

the right side

right side

Chalky, thin, off-white to

cream, simple with

occasional axial striae.

Chalky, often thick,

off-white, simple,

with occasional axial

mple

striae

* Characters of adult shells only

mens with protoconchs; to Jacky Van Goethem, Claudine

Claes and Diana Hoortman (IRSNB) for help in search-

ing literature and access to the collection of IRSNB; to

Ricky Zandali, Marsh Harbour, Abaco, Bahamas, for the

loan and gift of specimens, and to Carol Skoglund for

donating type material of the new species to the USNM

and LACM as well as lending comparative material of

the four species for study.

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+ catalogue: | ~9 (1841).

Vokes, E. H. 1988. Muricidae (Mollusca: Gastropoda) of the

Esmeraldas beds, northwestern Ecuador. Tulane Studies

in Geology and Paleontology 21: 1-50.

Vokes, E. H. 1989a. Neogene Paleontology in the northern Do-

minican Republic. 8. The family Muricidae (Mollusca:

Gastropoda). ). Bulletins of American Paleontology 97 (332):

5-94.

Vokes, E .H. 1989b. Muricidae (Mollusca: Gastropoda) of the

Angostura Formation, northwestern Ecuador. Tulane

os in Geology and Paleontology 22: 107-118.

Vokes, E. H. 1996. One last look at the Muricidae. American

Conchologist 24 (4): 4-6.

THE NAUTILUS 120(2):66-75, 2006

Page 66

New buccinoid gastropods from uppermost Cretaceous and

Paleocene strata of California and Baja California, Mexico

Richard L. Squires LouElla R. Saul

Department of Geological Sciences

Califormia State University,

Northridge, CA 91330-8266 USA

[email protected]

County

Invertebrate Paleontology Section

Natural History Museum of Los Angeles

900 Exposition Boulevard

Los Angeles, CA 90007 USA

lousaul@earthlink net

ABSTRACT

Two new genera and three new species of extinct buccinoid

gastropods are described and named from the Pacific slope of

North America. The buceinid? Ornopsis? dysis new species is

from uppermost Maastrichtian ( (uppermost Cretaceous) strata

in the Dip Creek area, San Luis Obispo County, west-central

California. The fasciolarine fasciolarid? Saxituberosa new genus

is comprised of the lineage Saxituberosa fons new species, from

lower Paleocene (Danian) strata in Los Angeles County, south-

em California, and Saxituberosa titan (Waring, 1917), from

middle Paleocene (Selandian) strata in Ventura and Los Ange-

les counties, southern California, and northern Baja California,

Mexico. The fusinine fasciolarid? Perrilliata califia new genus

and new species is known from middle Paleocene (Sel indian)

strata in Ventura County, southern California and northern

Baja California, Mexico. Ornopsis? dysis and Perrilliata califia

are similar in morphology and geologic age to Gulf Coast Or-

nopsis (Ornopsis) glenni Wade, 1916, and Fasciolaria? plum-

meri Gardner, 1933, respectively; whereas probable congeners

for Saxituberosa are unknown, and it appears to be endemic to

the northeastern Pacific.

INTRODUCTION

Four species, three of which are new, are described in

this report. They are of similar aspect in having fusiform

shells with tabulate whorls ornamented by ievolis and

spiral ribs. They are recognized as being of three differ-

ent genera, two of which are new.

Saul (19S6a), in her exploratory work on shallow-

marine mollusks from the Lake Nacimiento area, San

Luis Obispo County, west-central California, illustrated a

specimen of Ornopsis? n. sp. from the uppermost Cre-

taceous part of the El Piojo Formation along the east side

of Dip ¢ reek (Figure 1). Saul (1986a), furthe srmore, re-

ported that Ornopsis? n. sp. is apparently the “Trachytri-

fon” titan Waring, 1917, that was incladed in the Dip

Creek macrofaunal list provided by Taliaferro (1944).

This present work was initiated in order to nee a

fons new species, of

better understanding of the relationship between Ornop-

sis? 1. Sp. and “T.” titan. Saul’s Ornopsis? n. sp. is herein

assigned to Ornopsis? dysis new species, and its genus

assignment cannot be made with certainty until more

specimens are found. Ornopsis sensu stricto Wade, 1916,

heretofore has been reported with certainty only from

Upper Cretaceous (upper Campanian to upper Maas-

trichtian) strata of the southeastern United States (Sohl,

1964). “Trachytriton” titan is herein found to occur in

middle Paleocene strata in southern California. Its char-

acteristics separate it from the cymatiid Trachytriton

Meek, 1864, and “T.” titan is assigned to Saxituberosa

new genus, which is also represented by Saxituberosa

early Paleocene age.

The fourth species de scribed in this re port, Perrilliata

califia new genus and new species,

four. It occurs in middle Paleocene strata in northern

Baja California, Mexico.

The classification system used here follows that of

Bouchet et al. (2005). All of the new taxa are neogastro-

pods and are placed in superfamily Buccinoidea

Ralinesque, 1815. The familial cli assification for each of

the new taxa described below is tentative, especially be-

is the largest of the

cause no information is known about their protoconchs

nor about the anterior portions of their shells. Another

factor to take into account is the observation made by

Bandel (1993: 8), who stated that modern groups of gas-

tropods “can usually easily be connected with their rela-

tives that lived during Tertiary times, but when crossing

over into the Mesozoic comparisons become increasingly

difficult due to convergence observed in regard to the

teleoconch as well as the protoconch.” We agree with his

observation, and, furthermore, we believe that it is pos-

sible that the taxa described below might eventually be

placed in new suprageneric categories. This study, nev-

ertheless, adds substantially to our knowledge of “Pacific

slope of North America latest Cretaceous to Paleocene

neogastropods, a rather poorly known Eeour: Institu-

CAS, California

IGM, Mexico

tional abbreviations usecl in the text are:

Academy of Sciences, San Francisco;

R. L. Squires and L R. Saul, 2006

e 120

Dip Creek

Pinyon Ridge

4 Warm Springs

Mtn.

Figure 1. Location of formations bearing the new taxa.

Museo del Paleontologia del Instituto de Geologta;

LACMIP, Natural History Museum of Los Angeles

County, Invertebrate Paleontology Section; PU, Purdue

University; UCLA, University of California, Los Angeles

collections now housed at LACMIP): LSJU, Leland

Stanford Junior University (collections now housed at

CAS).

STRATIGRAPHY

The ages and depositional environments of all of the

formations bearing the new taxa discussed in this paper

can be found in the following papers: El Piojo Formation

Saul, 1986a; Seiders, 1986, 1989; Squires and Saul,

1993): lower San Fr incisquito Formation (Squires,

1997): lower Santa Susana Formation (Squires, 1997);

and Sepultura Formation (Squires, 1997).

PALEOBIOGEOGRAPHIC IMPLICATIONS

A thorough but not exhaustive search of the literature

revealed that Ornopsis? dysis and Perilliata califia are

most similar to New World gastropods from the Gulf

Coast of the United States, whereas Saxituberosa is ap-

parently endemic to the study area. Details of the mor-

phologic comparisons are given under “Systematic Pale-

ontology.

The latest Cretaceous Ornopsis? dysis is similar to

Ornopsis (Ornopsis) glenni Wade, 1916, the type species

of this genus. Ornopsis known with certainty (Sohl,

1964) ) only from upper Campanian and Maastrichtian

beds in Tennessee and Arkansas. Re ports of Ornopsis

from elsewhere in the world are highly doubtful (Sohl,

1964). As mentioned by Wade (1926), there might be an

occurrence of Ornopsis from Maastrichtian strata in the

Netherlands. This occurrence is based on a gastropod

reported by Kaunhowen (1897: 88-89, pl. 13, fig. 13) and

idenfied by him as Fusus (Hemifusus) ne reidiformis

Kaunhowen, 1897. It has the fine -spiral sculpture and

strong collabral structure like that of Ormopsis glenni, as

well as the characteristic single fold above the siphonal

canal. Reports of Ornopsis from the Congo basin and

Angola, west-central Africa (Rennie, 1929; Darteville

and Brebion, 1956) are highly doubtful because the

specimens are internal molds (Sohl, 1964), and one of

the specimens (Darteville and Brebion, 1956: pl. 6, figs.

5a, 5b) has an umbilicus, which is a feature not associated

with genus Ornopsis.

Based on the known distribution of Ornopsis, it is very

plausible that surface currents flowing westward from

the southeastern United States toward the Pacific slope

of North America allowed for the dispersal of this genus

into California. These currents, which existed during the

Late Cretaceous (Gordon, 1973; Johnson, 1999) and con-

tinued into the Paleocene and Eocene (Saul, 1986b:;

Squires, 1987), were part of a circumglobal-tropical cur-

rent that contributed to a widespread dispersal of marine

biota (Haq, 1981).

The Paleocene Perrilliata califia is most similar to Fas-

ciolaria? plummeri Gardner, 1933, and to Fasciolaria

new species Gardner, 1933, both from lower Paleocene

(Danian) strata of the Midw ay Group in Texas. It is plau-

sible that this genus was dispersed westward ae the

Age : New Taxa Ranges Turritella

asian ae Zones

g5*

? ? ?

dysis

Upper Up. Maas

Cret.

Figure 2.

Ages of stage boundaries from Gradstein et al. (2004). Turri-

tella zones from Saul (1983

Middle

Selandian

T. peninsularis

Paleocene

T. peninsularis

qualeyi

Lower Paleogene

T. peninsularis

adelaidana

Chronostratigraphiec positions of the new taxa

Page 68 THE NAUTILUS, Vol. 120, No. 2

‘ee 12

Figures 3-13. New fasciolariid? gastropods. Specimens coated with ammonium chloride. 3-7. Ornopsis? dysis new genus and species.

34. Paratype LACMIP 13352, LACMIP loc. 26527, height 31.1 mm, diameter 20.9 mm. 3. Apertural view. 4. Abapertural view. 5-7.

Holotype LACMIP 7564, LACMIP loc, 26525, height 37.6 mm, diameter 32 mm. 5. Apertural view. 6. Apertural view turned slightly to

right. 7. Abapertural view. 8-11. Saxituberosa fous new genus and species. 8-9. Holotype LACMIP 13354, LACMIP loc. 21581, height

55.6 mm, diameter 40.4 mm. 8. Apertural view. 9. Abapertural view. 10. Paratype LACMIP 13355, LACMIP loc. 215S0A, apertural/

right-lateral view (specimen crushed), height 48.4 mm, diameter 35 min. LL. Paratype LACMIP 13356, LACMIP loc. 1588, abapertural

view, height 52.5 mm, diameter 42 mm. 12-13. Saxituberosa titan (Waring, 1917) new combination, hypotype LACMIP 10983, LACMIP

loc, 22658, height 60.4 mm, diameter 36.2 mm. 12. Apertural view. 13. Right-lateral view

fe)

R. L. Squires and LR. Saul, 2006 Page 69

Figures 14-19. New fasciolariid? gastropods. Specimens coated with ammonium chloride. 14-15. Savituberosa titan (Waring,

1917) new combination. 14. Abapertural view of same specimen shown in Figures 12-13. 15. Hypotype LACMIP 10982, LACMIP

loc. 22701, slightly oblique right-lateral view, height 53.1 mm, diameter 28.8 mm. 16-19. Perrilliata califia new genus and species

16-18. Plasto-holotype IGM 4432, PU loc. 1334, height 101.3 mm, diameter 51.5 mm. 16. Apertural view. 17. Apertural view,

closeup of growth lines on inflated peripheral part of last whorl. 18. Abapertural view. 19. Paratype LACMIP 13357, LACMIP loc.

22330, abapertural view, height 74.5 mm, diameter 66.2 mm.

Page 70

THE NAUTILUS, Vol. 120, No. 2

CAGE

Figures 20-23.

21. Saxituberosa fons new genus and species, see Figure 11. 2

23. Perrilliata califia new genus and species.

Gulf Coast into California, via the same circumglobal-

tropical current system mentioned above.

SYSTEMATIC PALEONTOLOGY

Order Neogastropoda Thiele, 1929

Superfamily Buccinoidea Rafinesque, IS15

?Family Buccinidae Rafinesque, 1815

Discussion: Buccinids are of medium size, having a

fusiform shell in which the spire makes up 40 to 50% of

the total shell height. The smooth protoconch is pau-

cispiral (approximately two whorls) and is low. There is

usually no collar on the ramp. The shoulder usually has

tubercles and collabral ribs. A posterior “notch” and an

umbilicus can be present. The columella is callused and

almost alw ays smooth. The outer lip is smooth or with

small teeth; the interior of the outer lip is smooth or

lirate. The siphonal canal is short to moderately long, and

the siphonal 7 isciole is usually strong, twisted to the left,

and upturned. The operculum is chitinous. The crowth

line is generally prosocline on the ramp but oy Metoclne

elsewhere.

Genus Ornopsis? sensu stricto Wade, 1916

Discussion: Wade (1926) placed Ornopsis in family

Fusidae Swainson, 1840. According to Ponder and

Warén (1988), this family name, base d on homonymy, is

unavailable and is equivale nt to family Fasciolariidae

Gray, 1853. Wenz (1941) placed Ornopsis in the family

Buccinidae Rafinesque, 1815, but Soh! (1964) placed it

in the family Fasciolariidae. Snyder (2003: 24) included

Ornopsis in his list of “genera removed from family Fas-

iolariidae.” He made no taxonomic decisions in his

Growth-line trends of the new taxa, abi pega view. 20. Ornopsis” dysis new genus and species, see Figure 4.

22. Saxituberosa titan (Waring, 1917) new combination, see Figure 14.

work, relying instead on previously published work; nev-

ertheless, ihe. did not report who removed Ornopsis from

the fasciolariids nor the basis for this removal.

Bandel (1993: 40) reported that the relatively simple

embryonic whorls and smooth larval whorls of the pro-

toconch of Ornopsis Wade, 1916, probably indicate that

this genus is a buccinid. In addition, the low spire and

rounded last whorl of Ornopsis? dysis new species also

resemble that of a buccinid. This new species, however,

also has a single fold on the columella immediately pos-

terior to the siphonal canal. The only buccinid that we

know of that has this type of fold is Afe r Conrad, 1858, a

genus traditionally placed in the tudiclids, but placed by

Fraussen and Hadorn (1999) in the family Buccinidae.

The presence of a single fold on the columella immedi-

ately posterior to the siphonal canal, however, is not

unique to any one family, as it can be found in certain

members of other families including the following: sar-

ganids (e.g., Sargana Stephenson, 1923), but in compari-

son to O.> dysis these gastropods have umbilicate shells

with ornate sculpture: strepsidurids (e.g., nls eae

Swainson, 1540), but these gastropods have small shells

with a narrow spire and usually a smooth last whorl;

tudiclids (e.g., Tudicla Roding, 1798, and Rapopsis Saul,

1988S). but he se gastropods h vave shells with a very de-

pressed s spire, and, in the case of Rapopsis, the shell is

also umbilicate.

Although three subgenera of Ornopsis were recog-

nized by Soil | 1964), paced on shell shape and srowth-

line trend, the new species described below is more like

Ornopsis sensu stricto because the other two subgenera,

Ripleyella Harbison, 1945, and Pornosis Sohl, 1964, have

shells that are slimmer, more elongate, higher spired,

R. L. Squires and L R. Saul, 2006

and possess a sinuous growth-line trend with a strong

sinus. Ormopsis? dysis new species, is most like the buc-

cinid? Ornopsis (O.) glenni because the new species has

the following features: bucciniform shape, prominent

sculpture, posterior collar, a single strong fold above the

siphonal canal, a posterior siphonal notch, and a twisted

siphonal canal.

Type Species: Ornopsis (Ornopsis) glenni Wade,

1916, by origin: al de ‘signation; Late Cretaceous (late

Campanian and Maastrichtian) . Tennessee and Arkansas.

Ornopsis? dysis new species

(Figures 3-7, 20)

Ornopsis? n. sp. Saul, 1986a: 30, figs. 57-58.

Diagnosis: Large Ornopsis? with weak collar, wide

pleur: al angle, and strong columellar fold.

Description: Shell medium (up to 36 mm estimated

height and 21 mm diameter, same specimen). Juvenile

and early adult shell moderately inflated (height to di-

ameter ratio approximately 1.5, estimated); adult shell

more inflated (height to diameter ratio approximate sly

1.2). Buccinoid. Spire low, approximately 38% of f shell

height. Pleural angle approximately 90°. Protoconch and

upper spire anikcnewn: Teleoconch approximately three

whorls. Whorls inflated medially but constricted poste-

riorly on ramp. Ramp broad, slightly concave, usually

smoothish, and bearing weak a collar. Ramp

without spiral ribs or, on adult specimens, with two very

weak ribs. Suture moderately impressed, wavy. Sculpture

subdued and consisting of many collabral ribs intersect-

ing spiral ribs on inflated me dial part of whorls; inter-

sections demarked by weak nodes. Nodes most promi-

nent on shoulder and occasionally be ‘coming transversely

elongate toward outer lip. Collabral ribs moderate sly

widely spaced and most prominent on shoulder and on

inflated portion of last whorl; approximately 11 nodes on

shoulder of last whorl with nodes becoming stronger to-

ward outer lip. Collabral ribs usually extending very

weakly across ramp and obsolete on base and neck of last

whorl. Spiral sculpture consisting of numerous narrow

and moderately widely spaced ribs, more prominent than

collabral ribs. Approximate ly 1-13 spiral ribs on last

whorl from shoulder to neck; spiral ribs becoming

slightly weaker on neck. Aperture narrow on juvenile and

early adult shells but moderately wide on adult shell.

Posterior notch present. Columella slightly concave and

smooth except for single fold just above (posterior to)

siphonal canal where aperture becomes constricted to

form narrow siphonal canal, twisted to left. Growth lines

sinuous, prosocline on ramp, slightly opisthocline on pe-

riphery, and sigmoidal on base and neck area.

Holotype: LACMIP 7564, tip of spire and siphonal

canal missing, 37.6 mm height, 32 mm diameter.

Paratypes: LACMIP 133

Type Locality: LACMIP loc. 26525

52 and (unfigured) 13353.

Geologic Age: — Late Cretaceous (latest Maastrichtian).

Distribution: —E] Piojo Formation, Dip Creek, Lake

Nacimiento area, northern San Luis Obispo County,

west-central California.

Etymology: Greek dysis, meaning a dipping or setting;

in reference to Dip Creek.

Discussion: The new species is based on three speci-

mens. Their sculpture is subdued, probably because of

preservation. They range in estimated height from 2]

mm to 36 mm. None is complete.

Ornopsis? dysis is similar to Ornopsis (O.) glenni

Wade (1926: 463, pl. 24, fig. 1; Sohl, 1964: 215-216, pl.

29, figs. S-10, 15, 16), from Upper Cretaceous (Campa-

nian to Maastrichtian) strata in Tennessee and Arkansas

(Sohl, 1964), but the new species differs by being smaller

and having a narrower pleural angle (at least on the adult

shell), narrower ramp, stronger posterior collar, more

closely spaced and more spiral ribs, and stronger colla-

bral ribs.

Ornopsis? dysis resembles Hydrotribulus nodosus

Wade (1916: 465, pl. 24, figs. 4, 5; Wade, 1926: 147, pl.

51, figs. 6, 7; Sohl, 1964: 245-246, pl. 36, figs. 19, 20)

from Upper Cretaceous (Campanian to lower Maastrich-

tian) strata in Tennessee (Sohl, 1964), but the new spe-

cies differs by having rounded rather than tabulate shoul-

ders, a fold on the colamel la rather than a ridge, absence

of nearly cancellate sculpture, and absence of a strong

parieti ul ‘tooth,

The new species somewhat resembles Buccinopsis

crassa (Wade, 1917: 291, pl 19, figs. 6, 7; Wade, 1926:

145, pl. 50, figs. 9-12; Sohl, 1964: 189, pl. 22, figs. 1, 2)

from Upper Cretaceous (upper Campanian to lower

Maastrichtian) strata of Tennessee and Texas, but the

new species differs by having a much stronger fold on the

columella and an absence of a highly inelaed: broad

siphonal fasciole bordered above by a narrow deep slit.

Bandel (1993: 40) reported that Buc inopsis is probably

a buecinid.

?Family Fasciolariidae Gray, 1S53

Discussion: — Fasciolariids are of medium to large size,

having a fusiform shell in which the spire makes up 40 to

50% of the total shell height. The smooth protoconch is

usually moderately high (approximately three whorls),

but in rare cases, it is lew or bulbous. The teleoconch can

have spiral ribs or can be smooth. A collar is usually not

present. The shoulder bears tubercles; coll: tbral ribs

most prominent on the shoulder, wheras spiral ribs dom-

inant elsewhere. The columella has none or one to three

folds, extending posteriorly upward into aperture. The

outer lip edge is smooth or with small teeth. The outer

lip interior is smooth or lirate. The siphonal canal is long

and narrow, and the siphonal fasciole is usually weak,

slightly twisted to the left, and slightly upturned. The

growth line is generally prosocline to almost orthocline.

?Subfamily Fasciolariinae Gray, 1$53

Discussion: On fasciolariines, the shoulder bears tu-

bercles, the columella has one to three folds, and the

Page 72

THE NAUTILUS, Vol. 120, No. 2

ramp is concave. The fusiform shape, high spire, concave

ramp, two teeth on the columella, tabulate whorls shoul-

dered with tubercles, and well developed collabral and

spiral ribs make it likely that Saxituberosa is a fasciolari-

ine.

Genus Saxituberosa new genus

Type Species: Saxituberosa titan (Waring, 1917);

middle Paleocene, southem California.

Description: Shell moderately large (up to approxi-

mately 70 mm height). Fusiform-tabulate. Spire moder-

ately high, approximately 44 to 45% of shell height. Pleu-

ral angle approximate ‘ly 70°. Protoconch and uppe rmost

spire ‘Gaknown: Teleoconch up to at least 5.5 whorls.

Whorls inflated peripherally but constricted posteriorly

on ramp. Ramp broad, smoothish, slightly concave, and

bearing very weak subsutural collar. Sculpture consisting

of eallabral ‘ribs interse cting spiral ribs on inflated medial

part of whorls, intersections demarked by nodes (11-13

on last whorl), usually strong but dying out anterior and

posterior to shoulder. Spiral ribs prominent on anterior

part of last whorl. Aperture elliptical. Columella slightly

concave and bearing two oblique folds just posterior to

siphonal canal; folds near es side of columella. Si-

phon: il canal slightly twisted to left. Growth lines proso-

cline on ramp, slightly opisthocline to orthocline on pe-

riphery, sinuous on anterior part of most inflated part of

body whorl, and nearly orthocline on neck.

Geologic Age: Early Paleocene (Danian) to middle

ee ene (Selandian)

Combination of Latin saxum, meaning

Etymology:

rock or stone, and Latin tuberosus, meaning full of

lumps; in reference to the stony tubercules that help

characterize this genus.

Discussion: Saxituberosa resembles genus Lupira

Stephenson, 1941, which Sohl (1964) ) placed in the fam-

ily Xancidae Pilsbry, 1922. As far as it known, this genus

is restricted to U /pper Cretaceous ( upper Pam ge to

Maastrichtian) strata in the southeastern United States

(Sohl, 1964). Saxituberosa differs from ae by having

a higher spire, fewer folds (two) on columella (rather

than rarely two and usually three to six), more anteriorly

located folds, more widely spaced folds, fusiform shell

shape (rather than pyriform), narrower pleural angle, less

prominent spiral sculpture (especially near shoulder,

fewer spiral ribs on inflated medial part of last whorl,

(rather than 13-16),

tuberculate nodes on shoulder and ane of uniformity

in size of nodes on inflated medial part of last whorl, and

fewer nodes (11-13) on shoulder

inner lip not heavily callused.

Saxituberosa somewhat resembles the shape and

sculpture of various species of genus Taioma Finlay and

Marwick, 1937, but Taoima lacks folds on its columella

and has a growth line that trends opposite to the direc-

tion of Saxituberosa. The familial affinities of Taioma are

not well understood. Stilwell et al. (2004) reviewed the

history of the contentious placement of this genus, and

they concluded, with some reservation, that it a fusinine

fasciolariid. Taioma is apparently confined to the Creta-

ceous and Tertiary of South America, Antarctica, New

Zealand, and Greenland (Griffin and Hiinicken, 1994:

Stilwell et al., 2004), Bouchet et al. (2005) classified

Taioma as a neogastropod in its own family: family Taio-

midae Finlay and Marwick, 1937; superfamily unas-

signed.

The location of the two folds near the ventral side of

the columella and near the anterior end of the aperture

of Saxituberosa is very similar to that of Fasioplex Mar-

wick, 1934, known from the Eocene of New Zealand.

The new genus differs considerably from Fasioplex by

having a much higher spire, a fusiform rather than a

vasid-like shell, and a narrower aperture.

Saxituberosa fons new species

(Figures S—11, 21)

Diagnosis: A Saxituberosa with moderately low spire

and strongly noded sculpture. Pleural angle 90°. Suture

can be obscured by shell material on ramp, Columella

with two folds immediately posterior to siphonal canal.

Description: Shell medium large (up to 55 mm esti-

mated height and 47.7 mm diameter, same specimen);

height to diameter ratio approximately 1.2. Fusiform-

tabulate. Spire moderately low, approximately 44% of

shell height. Pleural angle approximately S0° (all speci-

mens evaded) : Proteconch and upper spire nian

Teleoconch with at least five whorls. Whorls inflated me-

dially but constricted posteriorly on ramp area. Ramp

broad, concave, smooth: possibly bearing very weak sub-

sutural collar. Ramp can be filled by mdnepiea: Suture

usually indistinct. Sculpture consisting of many collabral

ribs intersecting spiral ribs on inflated medial part of

whorls; intersections demarked by strong nodes or tu-

bercles. Collabral ribs moderately closely spaced and

most promine nt on shoulder and on inflated portion of

last whorl; approximately 12-13 nodes or tubercles on

shoulder of last whorl. Collabral ribs not present on ramp

and obsolete on base and neck of last whorl. Spiral sculp-

ture te of numerous strong and moderately

closely spaced ribs. Ramp without spiral ribs. Spiral ribs

most prominent on last whorl in area from shoulder to

neck. Spiral ribs unnoded on base of last whorl and on

neck. Aperture moderately wide. Columella with two

folds immediately posterior to siphonal canal where ap-

erture becomes constricted to form siphonal canal.

Growth lines sinuous, prosocline on ramp, opisthocline

on periphery, prosocline near neck area, and almost or-

thocline on neck.

Holotype: LACMIP 13354, siphonal canal missing,

55.6 mm height, 40.4 mm diameter.

LACMIP 1335

LACMIP loc. 1588.

Paratypes: 5 and 13356.

Type Lo ality:

Geologic Age: Early Paleocene (Danian) (Turritella

peninsularis qualeyi Zone of Saul, 1983).

R. L. Squires and L RB. Saul, 2006

Distribution: Lower San Francisquito Formation,

Warm Springs Mountain, northern Los Angeles County,

southern California.

Etymology: Latin fons, meaning spring; in re ference

to Warm Springs Mountain.

Discussion: A total of 14 specimens of this new spe-

cies were studied. The specimens range in size from 23.1

mm height and 21.8 mm diameter (same specimen) to

70.2 mm height. None of the specimens is complete, and

the largest one is broken in half, longitudinally. The most

complete specimen is 55 mm height and 47.7 mm diam-

eter, with a height to diameter ratio of 1.2. Preservation

of this species is poorer than the other two new species,

and the aperture and siphonal canal regions are crushed

and usually missing.

Saxituberosus fons is most similar to S. titan from

which it differs by having a wider pleural angle, lower

spire, much less ‘tabulate whorls, stronger tubeveules:

and occasionally broader ramp bordered by indistinct

sutures.

Saxituberosa titan (Waring, 1917) new combination

(Figures 12-16, 22)

Trachytriton titan Waring, 1917: 87, pl. 14, fig. 18

Penion titon |sic] (Waring).—Zinsmeister, 1953a: table 1, pl. 3,

figs. 22, 23.

Penion titan (Waring). Zinsmeister, 1974: 141-142, pl. 15, figs.

1, 2: 1983b: 1294, figs. 3H, 3 1.

not Penion cf. P. titan (Waring).—Paredes-Mejia, 1989: 256—

257, pl. 8, figs. 3, 4.

Diagnosis: A Saxituberosa with moderately high spire

and strong sculpture. Pleural angle approximately 70°.

Columella with two folds immediately posterior to sipho-

nal canal.

Description: Shell large (up to 67.4 mm height and

42.5 mm diameter, same specimen), height to diameter

ratio approximately 1.7. Subfusiform ar turreted spire.

Spire approximately 48% of shell height. Pleural angle

approximately 65°. Protoconch and upper spire un-

known. Teleoconch with at least 5.5 whorls. Whorls in-

flated medially but constricted posteriorly on ramp area.

Ramp broad, concave, smooth, and bearing very weak

subsutural collar. Suture moderately impressed, some-

what wavy. Sculpture consisting of spiral ribs inter secting

with collabral ribs, intersections demarked by strong

nodes or tubercles. Sculpture dying out above and below

inflated medial parts of whorls. Upper spire whorls with

single row of tubercles on tabulate shoulder; single row

gradually passing into double row of equally strong tu-

bercles on later whorls, Penultimate whorl shoulder ean

approximately 15 nodes. Last whorl with four spiral rows

of tubercles on most inflated part of whorl; poste rior pair

of rows closely to widely spaced and bearing strongest

tubercles. Last whorl shoulder with approximately 12-13

nodes. Anterior pair of rows on most inflated part of last

whorl with less projecting tubercles, becoming somewhat

elongated in parietal region. Anteriormost part of last

whorl with one to two rows of subdued nodes. Neck with

approximately six spiral ribs, unnoded. Aperture moder-

ately wide and elliptical. Columella straight to slightly

concave (on adults), with wide callused area, and bearing

two strongly raised folds immediately posterior to slightly

twisted?, narrow siphonal canal. Growth lines sinuous,

prosocline on ramp, opisthocline over periphery, sigmoi-

dal on base, and nearly orthocline on neck.

Holotype: CAS 61926.01 [= LSJU 142], very worn and

incomplete specimen (uppermost spire missing) with

columella buried in matrix, $1 mm height, 60.5 mim di-

ameter.

Hypotypes: LACMIP 10982 [= UCLA 59254] and

LACMIP 10983 [= UCLA 59253].

Type Locality: CAS loc. 61901.

Geologic Age: Middle Paleocene (Selandian) {= Twr-

ritella peninsularis Zone of Saul, 1983}.

Distribution: San Francisquito Formation, Pinyon

Ridge near Big Rock Creek, Valymero area, northern

Los Angeles County, southern California; and lower

Santa Susana Formation (“Martinez marine member” of

Nelson, 1925), Meier Canyon, Simi Hills, eastern Ven-

tura County, southern California.

Discussion: A total of 12 specimens of S. titan were

studied: five from the Pinyon Ridge area and seven from

the Simi Hills, including the two hypotypes UCLA 59253

and UCLA 59254, both illustrated by Zinsmeister (1974,

1983a, 1983b). Preservation of these specimens is gen-

erally good to excellent, except that each specimen is

missing the protoconch and most of the siphonal canal.

The dimensions of the specimens range from 31.6 mm

height and 18 mm diameter (same specimen) to 67.4 mm

height and 42.5 mm diameter (same specimen). The

smallest specimen is nearly complete and has a height to

diameter ratio of 1.75, whereas the largest specimen is

inissing most of its siphonal canal. Another, mostly com-

plete, specimen has 60.5 mm height and 36.1 mm diam-

eter, and its height to diameter ratio is 1.7. We conclude,

therefore, that the height to diameter ratio of S. titan is

approximately 1.7,

Waring (1917) placed this species in genus Trachytri-

ton Me ak 1864. The type species of f this monotypic ge-

nus is Buccinum vinculum Hall and Meek, 1854, and this

type species was reported by Wenz (1941) as being in the

ranellid genus Argobuccinum Bruguiére, 1792. Tra-

chytriton does not Ty we the falalate: whorls, subsutural

collar, strongly noded sculpture, nor the folds on the

columellar that characterize the shells studied herein.

Waring (1917) reported that titan is very similar to

Trachytriton tejonensis Gabb (1869: 154, pl. 26, fig. 34)

from the Eocene Tejon Group in Live Oak Canyon, The

holotype of T. tejonensis is mostly an internal mold, and

it has a varix, which is a morphologic feature not found

on titan, Stewart (1926) [1927] placed T. tejonensis into

synonymy with the ranellid of cquahia hornii (Gabb,

1864).

Zinsmeister (1974, 1983a, 1983b) assigned S. titan to

Page 74

THE NAUTILUS, Vol. 120, No. 2

the buecinid genus Penion Fischer, 1884, which ranges

from the early Paleocene (Danian) to Holocene (Wenz,

1941). Although Savituberosus and Penion can have simi-

lar shell shape, the presence of one or two columellar

folds on Saxituberosus readily distinguishes it from Pe-

nion. Zinsmeister (1974, 1983a, 1983b) did not report

the presence of any folds on the two specimens of S. titan

that he illustrated, but when the specimen shown here in

Figures 12-14 was carefully cleaned by the j eae author,

two columellar folds were observed (Figure 13). The

other specimen, which is shown here in Figure 15, is

missing the part of the columella that bears the folds.

The specime n (IGM 4431) that Paredes-Mejia (1989:

pl. 8, figs. 3, 4) identified as Penion cf. P. titan (Waring),

which is from the Sepultura Formation in Baja Califor-

nia, Mexico, is not Saxituberosa titan even though it has

sculpture similar to that found on S. titan, This Sepultura

Formation specimen is a turrid because its ramp has a

well-developed, deep symmetrical sinus that is so char-

acteristic of turrids.

Saxituberosa titan is most similar to S. fons new spe-

cies, and S. titan differs by having a narrower pleural

angle, higher spire, much more tabulate whorls, and a

more distinct suture,

Saxituberosa titan strongly resembles the shell shape

of the fossil * . mayi Hanna and Israelsky (1925:

45, pl. 7, fig. 12), known from beds transitional A the

Heath Forme =n near Quebrada Mancora, in the ex-

treme northwestern coastal region of Peru. The locality

description for this species is very imprecise, but it is

likely that the species occurs in the transitional beds be-

tween the Heath and Mancora formations, both of which

are early Miocene in age, according to Dunbar et al.

(1990). Saxituberosa titan differs from “Surcula” mayi by

having spiral ribs and two columellar folds. It is likely

that “S.” mari belongs to genus Taioma Finlay and Mar-

wick, 1937, which was decussed earlier.

Saxituberosa titan resembles the cassid Galeodea

(Taieria) klingeri Kiel and Bandel (2003: figs. 6.6-6.8)

from the Upper Cretaceous (middle Santonian-lower

Campanian) Umzamba Formation in South Africa. The

new species differs by having folds on the columella,

stronger collabral sculpture, ‘straighter columella, less

twiste a posterior portion of the siphonal canal, and an

absence of a small pseudo-umbilicus. In addition, S. titan

apparently lacks a posterior canal.

?Subfamily Fusininae Wrigley, 1927

Discussion: On fusinines, the shoulder is rounded, the

columella lacks folds, and shells can be large with high

spires and long siphonal canals. The subfamilial place-

ment of Perrilliata new genus is uncertain, mainly be-

cause the columella and aperture are not complete. In

addition, the growth lines on the ramp area are not pre-

served. Perrilliata might be a fusinine based on its fusi-

form shape, large size, high spire, and absence of any

columellar folds, but its strongly shouldered whorls with

tubercles, as well as strong spiral ribs overlying collabral

ribs, however, are features that are not usually found on

fusinines.

Perrilliata new genus

Type Species: Perrilliata califia new species; middle

Paleocene (Selandian), southern California and Baja

California, Mexico.

Description: — Shell large, up to 101 mm height. Fusi-

form with very spire high. Tabulate whorls. Ramp mod-

erately broad and concave. Nodes strong on shoulder.

Spiral ribs very prominent and closely spaced on periph-

ery and base. Neck smoothish. Columella long, straight,

and callused.

Geologic Age: Middle Paleocene (Selandian).

Etymology: Named for Maria del Carmen Perrilliat

(IGM), in recognition of her important contributions on

Cretaceous and Cenozoic mollusks of Mexico.

Discussion: The new genus somewhat resembles Her-

corhyncus Conrad, 1868, known with certainty only from

Upper Cretaceous (upper Campanian to Maastrichtian)

strata in Tennessee, Mississippi, Alabama, and Georgia

(Wade, 1926; Sohl, 1964), but the new genus differs by

having a higher spire, a narrower ple iva angle, growth

lines prosocyrt rather than opisthocline on medial part of

last whorl, and sculpture obsolete on neck. In addition,

the new genus apparently does not have an umbilical

chink opposite the posterior end of the siphonal canal,

but the presence of an umbilical chink on Hercorhyncus

is a variable feature. The new genus might have a con-

striction at posterior end of neck, like that present on

Hercorhyncus.

Sohl (1964) and Snyder (2003) placed Hercorhyncus

in the a fasciolariids, but Dead ‘| (1993: 40) con-

sidered the genus probably to be a buccinid, based on its

protoconch, eee is similar to that of the buccinid Or-

nopsis. Cossmann (1901: 73) considered Hercorhyncus

to be a subgenus of Streptosiphon Gill, 1S67, and Wenz

(1943: 1306) considered Streptosiphon to be a synonym

of Afer Conrad, 1858, hence making Hercorhyncus a

subgenus of Afer, which, as mentioned earlier, was

placed in family Buccinidae by Fraussen and Hadorn

(1999). Sohl (1964: 220), however, cited that the aper-

tural features, the higher spire, and the lack of any col-

umellar folds negates that Hercorhyncus belongs to ei-

ther Streptosiphon or Afe r.

The new genus resembles Saxituberosa by having

tabulate widals with strong sculpture but differs from the

type species of Santtuberosa by having a much higher

spire, much more sinuous growth lines on the ramp, no

posterior collar, stronger spiral ribs, straighter columella,

and no folds on the columella.

Perrillata califia new species

(Figures 16-19, 23)

?*Penion” 1. sp. Paredes-Mejia (1989: 257-259, pl. §, figs. 1, 2).

Description: Shell large (up to LOL mm height and

51.5 mm diameter, same specimen) _ shell he ioht to di-

R. L. Squires and L R. Saul, 2006

ameter ratio oo ately 2.0. Fusiform. Spire very

high, approximately 5 52% of shell height-. Pleural angle

approximately 73°. Protoconch unknown. Teleoconch up

to at least eight hauls. Whorls tabulate. Ramp broad,

concave, and usually smoothish with occasional spiral

ribs. Suture moderately impressed and wavy; usually in-

distinct. Sculpture consisting of collabral ribs and inter-

secting spiral ribs. Intersections of ribs on shoulder de-

marked by tubercles: intersections of ribs on periphery of

juvenile specimens forming cancellated sculpture pat-

tern. Collabral ribs weak and moderately widely spaced,

most prominent on shoulder, somewhat less prominent

on base of whorls and becoming obsolete anteriorly to-

ward base of last whorl; approximately 12 nodes (tu-

bercles on adults) on shoulder of last whorl. Collabral

ribs can extend across ramp but, if so, are usually poorly

developed. Collabral ribs obsolete on neck. S Spiral sculp-

ture consisting of numerous strong and moderately

closely spaced ribs. Upper spire whorls with one spiral

rib anterior to shoulder: middle spire whorls with two

spiral ribs anterior to shoulder; lower spire whorls with

three spiral ribs anterior to shoulder. Spiral ribs most

prominent on last whorl in area from shoulder to neck.

Posterior end of neck possibly delineated by constriction.

Neck smooth. Aperture probably small and narrow. Col-

umella straight and callused, with slightly raised, smooth

area along left margin. Growth line sigmoidal around

nodes on shoulder area with antispiral sinus on shoulder,

prosocyrt on periphery, and nearly orthocline on neck.

Holotype: IGM 4432, 101.3 mm height, 51.5 mm di-

ameter.

Paratype: LACMIP 13357

Type locality: PU loc. 1334.

Geologic Age: Middle Paleocene (Selandian).

Distribution: Lower Santa Susana Formation (“Mar-

tinez marine member” of Nelson, 1925), Meier Canyon,

Simi Hills, eastern Ventura County, southern California;

and Sepultura Formation, Mesa San Carlos, northern

Baja California, Mexico.

Etymology: Named for California.

Discussion: The new species is based on two speci-

mens. The holotype, which is from float material derived

from the Sepultura Formation at Mesa San Carlos, has

been crushed dorso-ventrally. It is a large specimen

(101.1 mm height, 47.6 mm diameter), ), whose height to

diameter ratio is 2.1, and it shows very good preservation

of the sculpture. The sculpture on the spire is very simi-

lar to that found on specimens of Saxituberosa titan less

than approximately 45 mm in height. On the remaining

whorls of this large specimen, spiral sculpture is very

strong on the anterior portion of the penultimate whorl

and on the base of the last whorl.

The paratype. which is from the Simi Hills, consists of

just the very large peripheral part (42 mm diameter) of

the last whorl of an adult specimen.

The geologic age of the specimens from the Sepultura

Formation is not ane n with certainty because the speci-

mens are float material. The ge ologic age of the speci-

men from the Simi Hills, ewes +r, is well constrained as

being middle Paleocene (Selandian) based on the asso-

ciated mollusks. Using the Simi Hills specimen as con-

trol, we infer that the Sepultura Formation specimens

are the same geologic age.

The new species is similar in shape, size, and sculpture

to Fasciolaria? plummeri Gardner (1933: 246-247, pl.

22. figs. 1-3) from the Paleocene eine nee of

hie Midway Group of Texas. Dockery (1986: fig. 1) cor-

related the Midway G rroup to the lower Paleocene (Da-

nian) and correlated the Kincaid Formation to the low-

ermost part of the Danian. Gardner (1933) reported that

F.? plummeri is unusually large (93 mm height) for a

Paleocene gastropod and that this species is one of the

few elements in the Midway fauna that is reminiscent of

the Cretaceous. The new ae cies differs from F.? plum-

meri by having a wider pleural angle, occasional pi

ribs on the 1 ramp; slightly wider, more closely spaced, and

more spiral ribs on The anterior swollen part of the last

whorl; no spiral ribs on the neck; and no hint of a sipho-

nal fasciole.

The new species is somewhat similar to Hercorhyncus

(Haplovoluta) triliratus Sohl (1964: 223-224, pl. 30, figs.

17-20, 23-24), which is known from upper Campanian

to upper Maastrichtian strata in Tennessee, Alabama,

and Georgia. The new species differs by having a nar-

rower pleural angle, higher spire, more w hore, ramp

spirally ribbed rather than smooth, and spiral ribs on

swollen part of the last whorl more numerous and more

closely spaced.

The new species resembles Saxituberosa titan but dif-

fers from S. titan by having a much higher spire, no folds

on the columella, much stronger spiral ribs, and, as far as

it can be ascertained, a straighter columella.

ACKNOWLEDGMENTS

Maria del Carmen Perrilliat kindly provided high-quality

replicas of IGM specimens that were collecte d and fig-

ured by Paredes-Mejia (1989). Lindsey T. Groves and

Steffen Kiel critiqued the manuscript.

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ington, D.C.: Smithsonian Institution and The National

Science Foundation, pt. 1, xiii + 626 pp.].

Wade, B. 1916. New genera and species of Gastropoda from

the Upper Cretaceous. Proceedings of the Philadelphia

Academy of Natural Sciences 68: 455-471.

Wade, B. 1917. New and little known Gastropoda from the

Upper Cretaceous of Tennessee. Proceedings of the Phila-

delphia Academy of Natural Sciences 69: 280-304.

Wade, B. 1926. The fauna of the Ripley Formation on Coon

Creek. Tennessee. U.S. Geological Survey Professional

Paper 137: 1-272

Waring. C. A. 1917. Stratigraphic and faunal relations of the

Martinez to the Chico and Tejon of southern California.

Proceedings of the California Academy of Sciences, Series

4, 7(4): 41-124.

Wenz, W. 1935-1944. Gastropoda. Teil 1: Allgemeiner Teil

und Prosobranchia. In: O. H. Schindewolf (ed.), Hand-

buch de Paliozoologie, Band 6. Berlin: Gebriider Born-

traeger, pp. 1-1639. [Reprinted 1960-1961].

Wrigley, A. G. 1927. Notes on English Eocene Mollusca with

descriptions of new species. Il. The Fusinidae. Proceed-

ings of the Malacological Society of London 17(5-6): 216—

249,

Zinsmeister, W. J. 1974. Paleocene biostratigraphy of the Simi

Hills, Ventura County, California. University of California,

Riverside, unpublishe -d Ph.D. dissertation, xii + 236 pp.

Zinsmeister, W. J. 1953a. Late Paleocene (“Martinez Provincial

Stage”) molluscan fauna from the Simi Hills, Ventura

County, California. In; R. L. Squires and M. V. Filewicz

(eds.), Cenozoic Geology of the Simi Valley Area, South-

ern California. Pacific Section, SEPM Fall Field Trip Vol-

ume and Guidebook, Book 35: pp. 61-70.

Zinsmeister, W. J. 1983b. New late Paleocene molluscs from

the Simi Hills, Ventura County, California. Journal of Pa-

leontology 57(6): 1252-1303.

APPENDIX

LOCALITIES CITED

Localities are LACMIP, unless othervise noted. All

quadrangle maps listed below are U.S. Geological Survey

maps.

CAS 6190. [= LSJU 4]. Exact location unknown, see Waring

(1917: fig. 3) for general location; Calabasas Quadrangle

(7.5 minute, 1952, photorevised 1967), north side of Cala-

basas Simi Hills, Ventura County, southem California.

Coll: C. A. Waring, summer 1910.

21579. Limey and sandy shale in canyon bottom, south side of

East Fork Fish Canyon, 1158 m east and 1630 m south of

northwest corner of Warm Springs Mountain Quadrangle

(7.5 minute, 1958), Los Angeles County, southem Califor-

nia. Lower Paleocene (Danian). San Francisquito Forma-

tion (lower part). Coll: R. W. Webb and E. H. Quayle,

June 5, 1941.

21580. Concretionary shale in canyon wall about 9 m above

stream bed, south side of East Fork of Fish Canyon, 1160

m east and 1463 m south of northwest corner of Warm

Springs Mountain Quadrangle (7.5 minute, 1955), Los An-

geles County, southern California. Lower Paleocene (Da-

nian). San Francisquito Formation (lower part). Coll.:

R. W. Webb and E. H, Quayle, June 5, 1941.

21588. Concretions in shale along elongate ridge crest, south

side of Warm Springs Canyon, 1463 m north and 792 m

west of Warm Springs Mountain, Warm Springs Mountain

Quadrangle (7.5 minute, 1958), Los Angeles County,

southern California. Lower Paleocene (Danian). San

Francisquito Formation (lower part). Coll: R. W. Webb

and E. H. Quayle, June 16, 1941.

22330. Beds cropping out on nose of spur on northwest side of

Meier Canyon, approximately 183 m north of second “n”

in Meier Canyon, Calabasas Caleb ih (7.5 minute,

1952, photorevised 1967), south side of Simi Valley, Simi

Page 78

THE NAUTILUS, Vol. 120, No. 2

Hills, Ventura County, southern California. Middle Pale-

ocene (Selandian). Santa Susana Formation (lower part).

Coll: W. P. Popenoe, April 3, 1946.

92688. Near summit of Simi Hills, 61 m southeast of hill 2150,

on ridge trending almost due south of hill 2151, 1753 m

east and 3079 m south of northwest corner of Calabasas

Quadrangle (7.5 minute, 1952, photorevised 1967), Ven-

tura County, southern California. Santa Susana Formation

(lower part). Middle Paleocene (Selandian). Coll.: J. H.

Fantozzi, circa June, 1951.

26525. Approximately 232 m elevation, poorly sorted conglom-

eratic sandstone exposed on east side of Dip Creek, south

side of Lake Nacimiento, 427 m south and 61 m west of

northeast corner of section 30, T. 25 S, R. 10 E, Lime

Mountain Quadrangle (7.5 minute, 1945), San Luis

Obispo County, west-central California. Upper Creta-

ceous (uppermost Maastrichtian). El Piojo Formation.

Coll: R. B. Saul and L. R. Saul, December 31, 1977.

26527. Approximately 225 m elevation, south side of Lake

Nacimiento, poorly sorted conglomeratic sandstone on

east side of narrows of Dip Creek, 777 m south and 304 m

west of northeast comer of section 30, T. 25 S, R. 10 E,

Lime Mountain Quadrangle (7.5 minute, 1948), San Luis

Obispo County, west-central California. Upper Creta-

ceous (uppermost Maastrichtian). El Piojo Formation.

Coll: R. B. Saul and L. R. Saul, December 31, 1977.

PU 1334. Float material from eastern part of Cajiloa Creek on

northwestern slope of Mesa San Carlos, Baja California,

Mexico, Middle Paleocene (Selandian). Sepultura Forma-

tion. Coll.: L. M. Paredes-Mejia, circa 1957.

THE NAUTILUS 120(2):79-S0O, 2006

Page 79

Authorship of the Ovulidae (Gastropoda) of the Zoology of the

Voyage of the SAMARANG

Richard E. Petit

S06 St. Charles Road

North Myrtle Beach, SC 29582

[email protected]

ABSTRACT

The Mollusca portion of the Zoology of the Samarang was

authored by Arthur Adams and Lovell Reeve. The new species

of Ovulidae therein, universally attributed to Adams and Reeve

in the modern literature, were described by G. B. Sowerby H,

who should be cited as author.

Additional Keywords: Adams and Reeve, G. B. Sowerby II

The Zoology of the Voyage of H.M.S. SAMARANG was

edited by Arthur Adams. It contains monogré iphs by vari-

ous authors, the Mollusca being eoasilhored by Arthur

Adams and Lovell Reeve. The Mollusca monograph was

published in 1548 and 1850 as detailed in the References

Cited herein. Species placed in the genus Ovulum. the

subject of this paper, are in the part published in 14S.

Adams and Reeve’s first grouping under the Gastero-

poda [sic] is the Convoluta. Although not given a rank

by them, the Convoluta is a Family in Reeve’s Tabula

Methodica (Reeve, 1841, 1842) consisting of the genera

Erato, Cypraea, Ovula, Terebellum and Conus. In the

discussion under the heading Convoluta the authors state

that “out of from eighty to a hundred species of Cones

collected during the voyage of the Samarang, only four

proved to be new, the greater number of those of recent

discovery being anticipate ed by Mr. Cuming during his

researches among the Philippine Islands and desea! ed

in the ‘Conchologia Iconica.” They continue with the

following two sentences (Adams and Reeve, 1848: 17)

“The genus Ovulum, not having been examined

since the publication of Mr. Sowerby’s ‘Species

Conchyliorum.,’ afforded a greater amount of nov-

elty. Mr. Sowerby, junr., being engaged in prepar-

inga monograph of this genus for the forthcoming

number of his ‘Thesaurus,’ it was thought desirab le

to place the specimens collected in his hands for

comparison, and we are indebted to him for the

descriptions and figures of eleven new species.”

The fact that the descriptions and figures were both by

Sowerby has as ed the attention of all later workers

except Sowerby and the Adams brothers. All of the new

_ s are listed and correctly attributed to Sowerby by

-and A. Adams (1854, 1: 270-272) under either Am-

en or Volva, the spelling of Ovula bullata being

corrupted to Amphiperas bullulata.

Under Article 50.1.1 of the Code (International Com-

mission on Zoological Nomenclature, 1999), these names

should be attributed to G. B. Sowerby I. In the Thesau-

rus these new Ovulium species are correctly attributed by

Sowerby to himself with reference to the SAMARANG fig-

ures,

In the introduction, where Owen is given credit for

having produced a section of the Mollusca, Sowerby is

not mentioned. There are other oddities about this work

and a detailed paper is in preparation.

It is surprising that these names have not been prop-

erly attributed. However, this has happened in a few

other works where subse squent workers have paid heed

only to parts of particular interest without reading intro-

ductory material. Even the venerable Sherborn attrib-

uted them to Adams and Reeve. Modem monographs of

the Ovulidae that would be expected to be thorough

enough to catch errors such as this one have yet to be

written. Schilder’s (1932) The living species of Amphi-

peratinae, which includes all of the species involved in

this discussion, cites references to only a few works in ab-

breviated form and even then some are inaccurately dated.

The seemingly most complete modem work is Cate’s

(1973) A systematic revision of the Recent Cypraeid fam-

ily Ovulidae. All of the species from the SAMARANG are

included in Cate’s work and all are attributed to Adams

and Reeve. Although Sowerby’s Thesaurus, where the

names are correctly attributed, is in Cate’s list of Litera-

ture Cited (inisdated as 1848), no entries from that work

are listed in the chresonymies of any of the SAMARANG

species. It must be me mtioned that a Sowe rby entry does

appear under “Phenacovolva acuminata (A, Adams and

Reeve, 1848).” That reference is:

1859 Ovula acuminata; Sowerby 2nd Tndex Brit.

Shells: 21, plt. 20, fig. 3.

The meaning of “21” is not clear as Sowerby’s work is no

ia: o of “2] tcl S | ki t

paginated but Plate 20, figure 3, is identifie od by Sowerby

as “Ovula acuminata Brug.” This European species

Page 80

THE NAUTILUS, Vol. 120, No. 2

was originally described as Bulla acuminata Bruguiere,

1792, onda is now usually placed in the genus Rhizorus in

the family Retusidae. This clearly has nothing to do with

the SAMARANG species.

In the “Monograph of the genus Ovulum” of his Con-

chologia Iconica, Reeve also atubuted the names to Ad-

ams and Reeve. Most recently, Alison Trew’s (1992)

Henry and Arthur Adams's new molluscan names attrib-

utes the SAMARANG ovulids to Adams and Reeve. In

short, as stated above, all authors except G. B. Sowerby

II and the Adams have misattributed these names. The

eleven new names are listed below in alphabetical order

by species name, Current generic placement of these

taxa is beyond the scope of ree paper.

Ovulum acuminatum G. B. Sowerby II in Adams and Reeve,

1848

Ovulum bulla G. B. Sowerby II in Adams and Reeve, 1548

Ovulum bullatum G. B. Sowerby IL in Adams and Reeve, 15458

Ovulum coarctatum G. B. Sowerby TH in Adams and Reeve,

1S48

Ovulum concinnum G. B. Sowerby IL in Adams and Reeve,

1S48

Ovulum dentatum G. B. Sowerby I in Adams and Reeve, 1545

[Preoceupied; invalid; renamed Prosimnia renovata lre-

dale, 1930].

Ovulum formosum G. B. Sowerby UH in Adams and Reeve, 1545

Ovulum gracile G. B. Sowerby II in Adams and Reeve, 1848

Ovulum subesulavuan G. B. Sowerby II in Adams and Reeve,

1545

Ovulum recurcum G. B. Sowerby ILin Adams and Reeve, 1545

Ovulum subreflexum G.B. Sowerby I in Adams and Reeve,

1S48

Thirteen species of Ovulidae are treated in the SAMA-

RANG, the eleven listed above being new. Of these

eleven, ten are still regarded as valid (Dr. Gary Rosen-

berg, personal communication, December 2005), which

is tier remarkable.

This writer would like to express his appreciation to

Dr. Gary Rosenberg for reading and commenting on a

draft of this paper and providing information about

ovulids.

LITERATURE CITED

Adams, A. and L. Reeve. 1848-50. Mollusca. In: A. Adams (ed.)

The Zoology of the voyage of H.M.S. Samarang; under the

command of Captain Sir Edward Belcher, C.B., F.R.A.S.,

F.G.S. during the years 1843-1846. Reeve, Benham, and

Reeve, London. x + 87 pp., pls. 1-24 [Pages 1-24, No-

vember 1848; 25-44, May 1550; 45-87, August 1850; pp:

i-x (by Adams only) and plates not basen Dates from

ses 1922: exi.|

Adams, H. and A. Adams. 1853-58. The genera of Recent

ae arranged according to their organization, ee an

van Voorst, London, 2 2 vols. ee 1-256, pls. 1-32 (1853);

957-484, 2, 1-92, pls. 33-72 (1854); 2, 93-284, pls. eae

(1855); 2, 385-40, pls. 97-112 (1856); 2, 413-540, pls.

113-128 (1857); 2, 541-660, pls. 129-138 (1858). |

Bruguiére, J. G. 1759- 99. Histoire Naturelle des Vers. Ency-

clopedié Méthodique. 1(1): i-xviii, 1-344, 1789; 1(2): 345—

757, 1792. [For complete collation of this work, see Even-

huis and Petit, 2003, Zootaxa 207: 1-4]

Cate, C. N. 1973. A systematic revision of the Recent cypr aeid

family Ovulid: ve (Mollusca: Gastropoda). The Veliger,

15( (Supplement): i-iii, 1-116, unnumbered plates.

International Commission on Zoological Nomenclature. 1999.

International Code of Zoological Nomenclature. Fourth

edition. International Trust for Zoological Nomenclature,

London, xxix + 306 pp.

Iredale, T. 1930, Queensland molluscan notes, no. 2. Memoires

of the Queensl: ind Museum, LO(1): 73-SS, “i 9.

Reeve, L. A. 1841-42. Conchologia eee or complete

system of conchology; in which the Le pades and Conchif-

erous Mollusca are described and classified ; according to

their natural organization and habits. Longman, Brown,

Green and Longman’s, London. 2 volumes.

Reeve, L. 1842. |Tabula ee Proceedings of the Zoo-

logical Society of London, 9: 72-76 (March); Annals and

Magazine of Natural History, 9(56): 145-152. (1 April)

Reeve, L. 1865. Monograph of thee genus Ovultwim. Conchologia

Iconica, 15: [unpaginated text], 14 pls. [Plate 2 dated

March 1862 in error; a paper on this entire work is in

preparation. |

Schilder, F. A. 1932. The living species of Amphiperatinae.

Proceedings of the Malacological Society of London,

20(1): 46-64, pls. 3-5. :

Sherborn, C. D. 1922-32. Index Animalium sive index nomi-

num quae ab A.D. MDCCLVIIL generibus et specibus

animalium imposita sunt. Sectio Secunda. A kalendris Tanu-

ariis, MDCCCI usque ad finem Decembris, MDCCCL.

British Museum (Natural History), London, exlvii + 7,056

+ 114 pp.

Sowerby, G. B., Il. 1849. Monograph of the genus Ovilum.

Thesaurus Conchyliorum 2: 467-454, pls. 99-101

Sowerby, G. B., II. 1859. Illustrated index of British shells.

Simpkin, Marshall, and Co., London, pp. i-xv, pls. 1-24

with unnumbered plate explanations.

Trew, A, 1992. Henry and Arthur Adams's new molluscan

names, National Museum of Wales, Cardiff, 63 pp.

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Volume 120, Number 3

September 22, 2006

ISSN 0028-1344

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CONTENTS

NAUTILUS

Volume 120, Number 3

September 22, 2006

ISSN 0028-1544

Bernard Landau

Carlos Marques da Silva

M. G. Harasewych

Josiah Strauss

Thomas J. DeVries

Lindsey T. Groves

K. J. Butkas

M. L. Ostrofsky

Eugene V. Coan

Richard E. Petit

The genus Scaphella (Gastropoda: Volutidae) in the Neogene of Europe and its

paleobiogeographical implications... 0... ee 8]

A new record of introduced Cerion (Gastropoda: Pulmonata: Cerionidae) in

southeastern Flonase ane dee ake OME Paw ete hae we dds awe daee a a 94

A new early Miocene Muracypraca Woodring, 1957 (Gastropoda: Cypraeidae)

from the Pisco Basin of southern Peru... 0.0.0. 101

The status of unionid and dreissenid mussels (Bivalvia) in northwestern

Pennsylvania inland lakes... 1... ee ee eee 106

Replacement names and type material: examples from Hertlein and Strong

(1940-1951) and Keen (1958) 2.0... eee 112

Book Review ...........

. Me

‘

= Se

‘

‘ '

THE NAUTILUS 120(3):S1-93, 2006

Page 8]

The genus Scaphella (Gastropoda: Volutidae) in the Neogene of

Europe and its paleobiogeographical implications

Bernard Landau

Lisbon University Geology Centre and

International Health Centres

Av. Infante Dom Henrique, Areias Sa0 Joao

8200 Albufeira, PORTUGAL

[email protected]

Carlos Marques da Silva

Departamento e Centro de Geologia

Faculdade de Ciéncias, Universidade

de Lisboa, C6, Campo Grande

1749-016 Lisboa, PORTUGAL

[email protected]

ABSTRACT

The genus Scaphella supposedly has a long geological history

on both sides of the Atlantic, extending back to the Paleocene.

However, there are differences in both shell morphology and

ecological preferences between the New and Old World rep-

resentatives. This paper traces the history of Scaphella in the

Atlantic. It is suggested that the group originated in the Cre-

taceous/Paleocene Tethys Sea, a genus such as Caricella dis-

persing during this time to the New World, and thence giving

rise in the Neogene to Scaphella. In the Old World the record

of Scaphella is uninterrupted from the Paleocene to the middle

Pliocene, after which the genus disappeared from the Eastern

Atlantic. The genus Scaphella is heterogeneous, the European

species differing in certain constant shell characteristics from

the New World species. Scaphella carlae new species is de-

scribed trom the lower-middle Pliocene Mediterranean of the

Estepona Basin (Spain) and a possible second new species is

discussed, but not named due to the poor material available,

from the lower Pliocene Atlantic of the

(Spain).

Guadalquivir Basin

INTRODUCTION

The subfamily Scaphellinae H. and A. Adams, 1858,

comprises three extant genera; Ampulla Roding, 1798,

Scaphella Swainson, 1832, and Volutifusus Conrad, 1863

rane 19SS). In the Recent fauna they occur in the

western Atlantic Ocean, including the southeastern

United States, Gulf of Mexico and Caribbean (Scaphella,

Volutifusus), and the eastern Atlantic, from southern

Portugal and Spain to the West African coasts of south-

ern Morocco and the Canaries (Ampulla). In the early

fossil record another genus, Caricella Conrad, 1835, pre-

dominates, being present from the Cretaceous in the

Tethyan record (Bandel, 2003) and in the New World

Paleocene to Oligocene (Dockery, 1977). Bandel (2003),

however, placed Caricella in a separate subl umily, Cari-

cellinae Dall, 1907. In this paper we deal with the Old

and New World taxa traditionally placed in the genus

Scaphella.

The gastropods of the genus Scaphella today live in

warm water, with a Recent subtropical to tropical distri-

bution restricted to the western Atlantic, from the coasts

of North Carolina (USA), southward through the Gulf of

Mexico to Yucatan (Weaver and du Pont, 1970), the Ca-

ribbean, up to Colombia (Clench, 1946; Poppe and Goto,

1992). The genus has an widespread geological record in

the Americas, with a few records in the Paleocene, an

extensive Neogene history, but is not recorded from the

Eocene or Oligocene.

In the Old World the Scaphellinae have an equally

long geological record extending back to the Paleocene

of the Nor th Sea Basin (Ravn, 1933). The group flour-

ished in the Miocene, extending its distribution into the

Atlantic and reaching the southern Atlantic coast of Ibe-

ria in the late Miocene (Pereira da Costa, 1866). In the

Pliocene the Scaphellinae were abundant in the North

Sea Basin (Marquet, 1997). Their range extended into

the Atlantic, as far as central western Iberia ( (Mondego

Basin) (Silva, 2001) and into the Mediterranean, where

they were restricted to the Alboran Sea (Estepona Ba-

sin), except for a single unconfirmed report from Algeria

(Lamothe and Dautzenberg, 1907).

The last European records for Scaphella are from the

upper Pliocene of the North Sea Basin. Today the

Scaphellinae are represented in the European faunas by

the monotypic genus Ampulla Roding, 1791. Ampulla

priamus (Gmelin, 1791) occurs from the southern coast

of Portugal (Nobre, 1935-40), or possibly from the

southwestern coast of Portugal, were it is rare (G,

Calado, pers. comm., 2006), south to the Canaries and

southern Morocco (Poppe and Goto, 1992).

Pliocene Atlantic and Mediterranean records of

Scaphella are scarce. Chavan and Coatman (1943) and

Brébion (1964) recorded §. lamberti from the Pliocene

of the Loire Basin. Silva (2001) listed S. lamberti from

the Atlantic Pliocene of the Mondego Basin and Zbys-

zewski (1943, 1959) from the Lower Tagus-Sado Basin of

central-western Portugal. There is only one unconfirmed

report by Lamothe and Dautzenberg (1907) of S. lam-

berti in the Mediterranean from the Place ne of Algeria

Recent work on the rich lower Pliocene de “posits of the

Page 82

THE NAUTILUS, Vol. 120, No. 3

Atlantic Guadalquivir Basin and Mediterranean Este-

pona Basin (southern Spain) revealed the presence of

Scaphella in both these basins.

FOSSIL-BEARING LOCALITIES

The material discussed herein originates from three dis-

tinct Iberian localities, situated, from northwest to south-

east, at:

1. Vale de Freixo, Pombal region, central-western Por-

tugal. Atlantic. Mondego Basin. The Pliocene Carnide

Sandstone Formation generally consists of fine mica-

ceous sand without macro somatofossils ( (body fossils).

Locally, the lowermost section of this formation contains

a thin fossiliferous sequence consisting of a basal con-

glomerate and sand rich in fossil shallow marine mollus-

can shells. At Vale de Freixo, the basal fossiliferous beds

of the Carnide Sandstone have a maximum thickness of

approximately 1 m. These are dated as lower to middle

Pliocene, uppermost Zanclean to lowermost Piacenzian

(Silva et al., 2000; Silva, 2001; Silva et al., 2006).

2. Lucena, Huelva region, southern Spain. Atlantic.

Guadalquivir Basin. The sandy, near-shore deposits of

Lucena are part of the Arenas de Huelva Formation and

dated as lower Pliocene, Zanclean (Civis et al., 1987).

3. Velerin, Estepona region, southern Spain. Mediter-

ranean. Estepona Basin. These deposits consist of a va-

riety of different lithologies, from fine clayey sands (Vel-

erin carretera outcrop), deposited at relatively greater

depths, to coarse conglomerates (Velerin conglomerates

outcrop). These conglomerates, which must have been

deposited relatively rapidly or in storm conditions (Sanz

de Galdeano and Lopez Garrido, 1991), contain the rich-

est fauna, a curious mixture of large and small abraded

and perfectly preserved shells. These deposits are dated

as middle Pliocene, lower Piacenzian (Guerra-Merchan

et al., 2002).

For detailed location maps and a geological and strati-

graphical overview of the fossiliferous deposits covered

in this paper see La Pema et al. (2003) and Dell’ Angelo

and Silva (2003), for the Mondego Basin, Civis et al.

(1987) for the Guadalquivir Basin, and Sanz de Galdeano

and Lopez Garrido (1991) and Guerra-Merchén et al.

(2002), for the Estepona Basin.

The Pliocene malacofauna from all three of these ba-

sins, according to Silva (2001), La Perna et al. (2003), and

Landau et al. (2003), corresponds to the Mediterranean

Pliocene Molluscan Unit 1 (MPMU1) of Monegatti and

Raffi (2001).

The material herein discussed is housed in the follow-

ing collections: IRScNB: Institut royal des Sciences na-

turelles de Belgique; BLP coll.; B. Landau collection;

CMS coll.: Carlos Marques da Silva collection, Departa-

mento de Geologia da Faculdade de Ciéncias de Lisboa;

RM coll: collection R. Marquet (will be incorporated

into the collection Institut royal des Sciences naturelles

de Belgique in the near future); CG coll. = Chris Garvie

collection, USA; M-V coll.; Manuel Molin/Daniela Velo

collection, Bonares, Spain; AC coll.: Alain Cluzaud col-

lection, France.

SYSTEMATIC PALEONTOLOGY

The genus Scaphella

The type species of the genus Scaphella is Voluta junonia

Shaw, 1808, by subsequent designation of Gray, 1847

(Clench, 1946). Scaphella junonia is a western Atlantic

species, characterized by axially ribbed post-nuclear

whorls, prominent columellar plications and a well de-

veloped siphonal fasciole.

Clench (1946) recognized Scaphella sensu stricto char-

acterized by shells that were a little more massive,

nuclear whorls extended and the calcarella usually worn

away; and Scaphella (Aurinia) H. and A. Adams, 1853, in

which the shell is strong, but not massive, and the

nuclear whorls have a str ongly developed calcarella. In a

more recent work (Weaver aad du Pont, 1970), this di-

vision into subgenera was confirmed by differences in

the radular structure. According to Clench (1946), the

typical subgenus has wishbone shaped rachidian teeth,

with the central denticle rather long and narrow, and the

lateral shanks extending behind rather than to the sides,

but no lateral denticles. In the subgenus Aurinia the

rachidian teeth have a very strong central denticle, with

shanks above extending at ‘almost right angles, and at the

base two very small iateral teeth, fused to the central

denticle (Clench, 1946).

Later, Weaver and du Pont (1970) recognized three

subgenera. The subgenus Scaphella sensu stricto was

characterized bya papilliform ao usually with a

spur-like calcarella present, the teleoconch “. . . is sculp-

tured with spiral{sic] lirae and incised lines ... ”, the

columella has three or more anterior plaits, and a sipho-

nal notch and fasciole are present (Weaver and du Pont,

1970: 140). The subgenus Aurinia differs in having a

larger protoconch, w ith a projecting calcarella, in lacking

a siphonal notch and fasciole, and in having no co lemele

lar plaits or only weak ones. They recognized a third

subgenus, Clenc hina Pilsbry aie Olsson, 1953, which dif-

fers from the nominal subgenus by smaller size, less solid

shell, usually with an attenuate od spire. These shell dif-

ferences are small, and both Pilsbry and Olsson (1953)

and Weaver and du Pont (1970) separated the subgenera

mainly by radular morphology. The nominal subgenus is

characterized by uniserial Y- -shaped radular te Ge with-

out side cusps, Clenchina has small Y-s -shaped teeth with

minute side cusps, and Aurinia has tricuspid rachidian

teeth, with the central cusp broadest and the lateral

cusps strong and sickle-shaped (Weaver and Du Pont,

mene

Poppe and Goto (1992) placed ‘shell similarity’ above

radular structure. They stressed that these subgener ra

were based mainly on radular characters, with ee

logically similar species having quite different radular

structure (Bayer, L971), and recognized only a single

genus Scaphe Ma, ethout subgenera.

B. Landau and C. M. da Silva, 2006

Page 83

In the Recent fauna the representatives of the genus

Scaphella are restricted to the western Atlantic. The

number of species recognized varies considerably, from

four (Weaver and du Pont, 1970) to 11 (Poppe and Goto,

1992) or 12 (Clench, 1946). All the Recent species, ex-

cept S. gouldiana (Dall, 1887)

squarish red or black spots in spiral rows on a paler

background, which has spiral bands in some species, and

most species have axial sculpture on the early teleoconch

whorls.

Scaphella in the fossil record

New WORLD

Scaphella is well ca dan a in the Neogene Caribbean

fossil record. It is found in the upper Miocene Gurabo

Formation of the Dominican Republic (Vokes, 1998) and

represented by possibly numerous species in the Florid-

ian Plio-Pleistocene (Petuch, 1994). It has not been re-

corded from the Tropical American Pacific. The earliest

representative of the Neogene group of Scaphella in the

American fossil record is the upper Miocene Dominican

Republic species S. striata (Gabb, 1873), which is most

similar to S. gouldiana in oe strong axial nodes at the

shoulder. Vokes (1998) noted that fie shells of S. striata

do not show the characteristic color pattern and were

probably monochrome. All the specimens from the

Florida Plio-Pleistocene illustrated by Petuch (1994)

have axial sculpture on the early teleoconch whorls and a

spotted color pattern.

The history of the genus in the New World before the

Neogene is far more complex. Dall (1907) suggested that

the Neogene group of Scaphella evolved in ihe western

Atlantic from the genus Caricella Conrad, 1835, which is

widespread in the Eocene and Oligocene western Atlan-

tic assemblages (Gardner, 1937).

Caricella shares a similar shell shape and color pattern

of squarish red dots (Dockery, 1977; MacNeil and Dock-

ery, 1984). It therefore seems that this color pattern is a

conservative character of the Caricella-Scaphella line of

volutes (MacNeil and Dockery, 1984). Gardner (1937)

described Caricella (Atraktus) pycnopecta Gardner,

1937 from the Shoal River Formation of northern

Florida and extended the upper range of Caricella into

the lower Miocene. In our opinion fis new taxon may

have been based on a juvenile specimen of Scaphella.

The holotype is 27.0 mm in height. The juvenile speci-

men of Scaphella sp. from the Flovidian Pliocene herein

figured (Figures 2-3) shows the same reticulate sculp-

ture, which later disappears in the adult stage. Unfortu-

nately we have not examined or been able to trace any

larger Scaphella specimen from the Miocene of Florida,

where they seem to be very rare.

Two species present in the Paleocene of the eastern

United States (Porters Creek Formation, Matthews

Landing Marl Member, Alabama) are much more similar

in their tall, elongated shape to the Scaphella represen-

tatives in the Old World. We refer to the former two

, have a color pattern of

species as the Paleocene New World eaters group.

The shell of Scaphella showalteri (Aldrich, 1886) (Fig-

ures 5-6) is very similar to the shells of the European

species, but does have faint axial sculpture on the early

teleoconch whorls, whereas the specimens of Caricella

leana Dall, 1890, (Figures 7-8) are very elongate, similar

to the shell of North Sea Basin Miocene Scaphella speci-

mens and most unlike any of the American Eocene-

Oligocene Caricella species.

OLD WoRLD

Traditionally, all the European Scaphella species had

been assigned to the genus Scaphella sensu stricto

Swainson, 1832. Darragh (1988) placed the European

species in the subgenus Aurinia H. and A. Adams, 1853.

However, as already noted by Darragh (1988) and Mar-

quet (1997), this allocation is problematic. There are cer-

tain shell features the Old World Scaphella species share

that are absent in the New World Neogene species. This

will be fully discussed further on.

The earliest records of the genus Scaphella in Europe

are Scaphella crenistria (Von Koenen, 1885) and

Scaphella faxensis (Ravn, 1902) from the Paleocene of

Denmark (Ravn, 1933). The shells of these Paleocene

species have the typical fusiform shape, but only three

columellar folds (Ravn, 1933) rather than the four or five

ones present in the shells of Neogene and Recent spe-

cies. Scaphella wetherellii Sowerby, 1836, from the lower

Eocene London Clay shows the typical Scaphella shape

ang also has three columellar folds (Edwards, 1855, pl.

23, fig. 4.). Scaphella honi Glibert, 1938 from the upper

Eocene, Bartonian (Wemmelian) of Belgium is some-

what unusual, with a carina on the last whorl bom

large sub-obsolete tubercles (Glibert, 1938, pl. 4, fig. 2),

bak the protoconch and early teleoconch whorls are simi-

lar to those of other European specimens, and like the

other early Scaphella, it has only three columellar folds.

Scaphella is then represented in the North Sea Basin

continuously from the Lower Oligocene, Lattorfian, of

Germany, by S . siemmsenii (Boll. 1851), which has an

elongate shell with a tall spire, no axial sculpture at all,

and no siphonal fasciole (Beets, 1950), to the Kruisschans

Sands (Marquet, 1997), Piacenzian, middle Pliocene

(Van Vliet-Lanoé et al., 2002). In the late Oligocene

Scaphella reached the Atlantic, where it was represented

by an undescribed species (Figures 11-13) from the up-

per Oligocene, Chattian of St-Paul-lés-Dax, Aquitaine

Basin, France (AC coll.).

In the Miocene Scaphella is represented in the North

Sea Basin by Scaphella bolli (Koch, 1861) (Figures 14—

17), and in the Atlantic by S. miocaenica (Fischer and

Tournouér, 1879) in the middle Miocene, Serravalian, of

the Loire Basin and S. tarbelliana (Grateloup, 1540) in

the lower Miocene, Burdigalian of the Aquitaine Basin

eee 1928). The soulhocnuast Miocene record is that

of Voluta lamberti Sowerby, 1816 in the upper Miocene,

aes. of southern Portugal, Algarve Basin (Pereira

da Costa, 1866).

20, No. :

NAUTILUS, Vol.

B. Landau and C. M. da Silva, 2006

In the Pliocene, Scaphella lamberti (Sowerby, 1816)

(Figures 19-22) flourished in the North Sea Basin and

was abundant in the Oorderen Sands of Belgium (Mar-

quet, 1997). In the Atlantic it is recorded from the Loire

Basin, northwestern France (Chavan and Coatman,

1943). Scaphella is also recorded from central-western

Portugal, Mondego Basin (Silva, 2001) and lower Tagus-

Sado Basin (Zbyszewski, 1943, 1959), southern Atlantic

coast of Spain, Guadalquivir Basin (BLP coll.) and the

Alboran Sea, Estepona Basin (BLP coll.), Lamothe and

Dautzenberg (1907) recorded S$. lamberti from the

Pliocene of Algeria, although we have not found any

further record to confirm this.

The origin of Scaphella

The subgeneric assignment of the European fossil

Scaphella species is problematic. Darragh (1988) placed

the European Cenozoic Scaphella species in the subge-

nus Aurinia, which according to him is characterized ‘by

the absence or reduction of the columellar plicae and the

absence of a siphonal fasciole.

Marquet (1997) already noted that Pliocene North Sea

Basin specimens of Sc -caphella lamberti do not fit neatly

within these characters, having strongly developed col-

umellar plications. The same can be said for the rest of

the European Scaphella species, which all have strongly

developed columellar plications.

One species, Scaphella (Aurinia) johannae Darragh,

1988, occurs in the Australian Aldingan, upper Eocene.

It is similar in shape to the European species and has no

sculpture on the early teleoconch whorls. As noted by

Darragh (1958, p. 21 7), that species is more akin to the

European stock of Scaphella than to the New World

Scaphella species, and could be regarded as an example

of a Tethyan element in the Australian Eocene fauna.

Thus, the shell characters of the Old World fossil spe-

cies of Scaphella do not fit into any of the three existing

subgenera of the genus. The early teleoconch whorls

have no axial sculpture; the aperture combines strong

columellar folds with the absence of a fasciole, and no

color pattern whatsoever has been observed in any speci-

men under either normal or UV light.

Bondarev (1997) discussed the biogeography and his-

tory of the ek Scaphellinae, stating that they orig-

inated in the Te thys, and placed their roots in the Me-

sozoic. Bandel (2003) supported the Mesozoic Tethyan

origin of the Volutidae with the description of a Creta-

ceous volutid assemblage from Egypt, from the southern

shelf area of the Tethys Ocean, on the African continent.

He also described the first and oldest member of the

genus Caricella from this upper Cretaceous Tethyan as-

semblage; Caricella (Misrimelo) klitzschi Bandel, 2003.

The origin of Scophella is therefore even less clear.

Any hypothesis on dispersal of the Scaphellinae must

encompass: the first appearance of Caricella in the

Tethyan Realm; the presence of Caricella and Scaphella

in the New World Paleocene, Caricella only in the New

World Eocene to Oligocene, Scaphella only in the New

World Miocene to Recent; the ie Ses: presence of

Scaphella in the Old World from the Paleocene to

Pliocene, but not of Caricella.

It is plausible to assume, as suggested by Dall (1907),

that the Neogene New World Gi caphella group evolved

from a Carice fla like ancestor, which migrated westward

from the Cretaceous Tethys into the proto- Caribbean

Sea along the predominantly westward flow of ocean

currents, ‘ong before the closure of the Central Ameri-

can Seaway, ih a more-or-less continuous tropical sea

existed at low latitudes (Vermeij and Rosenberg, 1993).

Based on the new data of Bandel (2003), it is however

more likely that both Caricella and Scaphella originated

in the Tethys Sea, and that both independently dispersed

to the New World, which could explain their presence

there in the Pale aoe and the similarity between the

Paleocene New and Old World Scaphella species. Sub-

sequently, as there is no record of any Scaplella in the

New World in the Eocene and Oligoce ne, he Neogene

New World Scaphella evolved from the New World Ca-

ricella, explaining the differences between this stock of

Scaphella species and the European stock, which re-

mained more closely similar to the original Tethyan

Scaphella.

However, none of the Recent Volutidae produce

planktonic larvae that could favor this westward transat-

lantic dispersal (Bouchet and Poppe, 1988; Darragh and

Ponder, 1998). Some Cenozoic Athleta species produced

planktonic larvae (Hansen, 1978); possibly some Lyria

species (Bouchet and Poppe, 1988); some Calliotectum

species (Bouchet and Poppe, 1995); and Provocator,

which appeared in New Zealand late in Pliocene time

(Maxwell, 2003). However, we must note that the Ath-

Figures 1-18.

Scaphella species. 1. Scaphella (Scaphella) junonia (Shaw, 1808) (BLP coll.). North New River Canal, South Bay,

Palm Beach County, Florida, USA, Bermont Formation, Pleistocene, height 79.6 mim, 2-3. Scaphella sp., juvenile (BLP coll.). APAC

Pit. Sarasota, Sarasota County, Florida, USA, lower Pliocene, Pinecrest Beds, height 32

2.1 mm. 4. Scaphella (Scaphella) martinshugari

Petuch, 1994 (BLP coll.). Early Pliocene, Pinecrest Beds Unit 10, Quality Aggreg gates Phase 6, Sarasota, Sarasota County, Florida.

Detail to show ratchet-shaped columellar folds. 5-6. Scaphella showalteri (Aldrich, 1886), (CG coll.) . Dixon’s Creek, Alabama River,

Wilcox County, Alabama , Porters Creek Formation, Matthews Landing Marl Member, Paleocene, height 22.8 mm. 7-8. Scaphella

leana (Dall, 1890), (CG coll.). Dixon’s Creek, Alabama River, Wilcox County, Alabama, Porters Creek Formation, Matthews Landing

Marl Member, Paleocene, height 26.9 mm. 9-10. Scaphella siemmsenii (Boll, 1851), juvenile

Chattian, upper Oligocene, height 19.3 mm. 11-13. Scaphella sp. (A

Oligocene, height 67.0 mm (photo A. Cluzaud). 14-17. Scaphella bolli (Koch, 1862)

(BLP coll.). Krefeld, Germany,

C coll.), Estoti, St.-Paul-lés-Dax, Landes, Chattian, upper

(BLP coll.). Borgerhout Ring Highway,

Antwerp, Belgium, Beechem Formation, Antwerp Sands, middle Miocene, height $0.4 mm. 18. Scaphella miocaenica (Fischer and

Tournouér. 1879) (RM coll.). Mathelin, Loire Basin, France, Serravallian, middle Miocene. Height 102 mm (photo Robert Marquet)

0, No.

THE NAUTILUS, Vol. 1

Page 86

B. Landau and C. M. da Silva, 2006

Page 87

letidae were considered as an independent family from

the Volutidae by Riedel (2000) (but as a subfamily of the

Volutidae by Bouchet and Rocroi [2005]).

In fact Bondarev (1997) stated that volutids are char-

acterized by their high level of provincialism. Bouchet

and Poppe (1988. p: 30) pointed out that “Volutes are the

most holobenthic of all ala ide *> and Darragh

(1988) noted that the volute fauna of the areas where

species occur today was largely established by mid-

Tertiary time, little having been added since. The fact

that, as far as we know, Scaplella in the New World

never dispersed into the Pacific during the Neogene, and

in the Old World it did not disperse into the Meditexra:

nean (except the Alboran Sea) or Paratethys, is a testa-

ment to its poor dispersal ability. This has not, however,

prevented the dispersal of some volutids over long dis-

tances, such as representatives of Alcithoe and Lyria,

that probably “hopped” along underwater bridges or

chains; these may have a short-lived demersal free-

swimming larval stage (Bouchet and Poppe, 1988S).

Bandel (2003) suggested that the protoconch-type of

the Cretaceous Egyptian Volutidae (including a Cari-

cella-like species) suggested a non- planktotrophic devel-

opment and that they ‘spread only by crawling young and

not by free-swimming larvae.

If Bondarev (1997) and Bandel (2003) are correct in

their hypothesis that the Scaphellinae originated in the

Cretaceous of Tethys Sea, and Dall (1907) is correct in

his hypothesis that Scaphella originated from Caricella, it

is likely that the New World Paleocene Scaphella sho-

walteri evolved from a Cretaceous/Paleocene Old World

ancestor, which emigrated to the New World from the

Tethys and in turn gave rise to the New World Scaphella

stock. The el age species traditionally assigned to

Scaphella, showing consistently distinct conchological

charactedsuies. therefore would constitute a separate

stock from those in the New World fauna, and might

even warrant a distinct genus-level taxon.

Family Volutidae Rafinesque, 1815

Subfamily Scaphellinae Gray, 1857

Genus Scaphella Swainson, 1832

Type species: Voluta junonia Shaw, 1808, by subse-

quent designation, Gray, 1847 (Clench, 1946). Recent,

western Atlantic Ocean.

Discussion: Whilst including the European species

traditionally assigned to Scaphella within this genus, we

stress that they differ from Scaphella sensu stricto and

the subgenus Clenchina in not having axial sculpture on

the early teleoconch whorls, and in having no siphonal

fasciole and no color pattern, and from the subgenus

Aurinia by again not having axial sculpture or color pat-

tern and having well developed columellar folds. Al-

though the number of columellar folds is similar in

Scaphella and its subgenera to that in the European

Cenozoic species, and in both the folds become more

oblique abapically, there is a subtle difference in their

shape. The folds in New World Scaphella specimens are

highly asymmetrical; the anterior face is much less steep

than the’ posterior face, giving the folds a ratcheted ap-

pearance (Figure 4). This is not true in the European

shells, in which the folds are elevated and syminetrically

rounded. This ratchet-like character of the columellar

folds of the New World Scaphella shells is not present in

Caricella, in which the folds are symmetrical and much

finer than in Scaphella.

Although the absence of color pattern in fossil shells is

not always a reliable guide to the color of the living

animals, almost all fossil Sc. aphella species from the New

World (except S. striata) are characterized by shells with

persistent color pattern (see Olsson and Petit, 1964:

Campbell, 1993: Petuch, 1994). Many other shells from

the Estepona and Mondego deposits have the color pat-

tern preserved and one would have expected some of the

strong spotted pattern of Scaphella to be seen if present,

but none has been observed in the specimens from Es-

tepona, Mondego, or in any other European fossil

Scaphella species.

Unlike Recent American Scaphella species, which are

tropical to subtropical, European fossil Scaphella species

were predominantly warm-temperate to subtropical.

Two of the three more southern deposits where Euro-

pean Scaphella have been found (Mondego and Este-

pona Basins), which were al aa tropical re-

spectively (see Silva and Landau, In press), reflect areas

where there was some degree of upwelling of cooler

nutrient-rich waters ( faridan et. al, 2004; Silva et al.,

2006). This ability of ae of temperate waters to

survive in a tropical zone subjected to the periodical

upwelling of colder nutrient rich waters has already been

observ ig for species of the genus Amalda ( Landau and

Silva, 2006). Even though European Scaphella were

widespread and diverse during the early Pliocene, the

genus did not survive subsequent Plio-Pleistocene cool-

ing, and the youngest record is of $. lamberti from the

upper Pliocene rutsechans Sands of Belgium, where it

is rare (Marquet, 1997). Interestingly, European

Scaphella did not follow the prevalent trend of south-

ward migration seen in many other gastropod taxa as a

Figures 19-31.

Scaphella species. 19-21. Scaphella lamberti (Sowerby, 1516),

(BLP coll.). ‘Broad form’, Vrasenedok, Kallo,

Onc V laande eren, Antwerp, Belgium, Oorderen Sands, middle Pliocene, height 133.0 mm. 22. Sc aphella lamberti (Sowerby, 1516),

BLP coll.).

Narrow form’, Vrasenedok, Kallo, Oost-Vlaanderen, Antwerp, Belgium, Oorderen Sands, middle Pliocene, height 176.0

mm. 23-26. Scaphella carlae new species, holotype, IRScCNB IST 6994, Ve lerin ¢ Jonglomerates, Velerin, Estepona, Spain, lower

Piacenzian, middle Pliocene, height 147.0 mm. 26. Detail showing rounded columellar folds. 27-29. Scaphella carlae new species.

Paratype (BLP coll.), Velerin Conglomerates, Velerin, Estepona, Spain, lower Piacenzian, middle Pliocene. Height 135.0 mm. 30-31.

Scaphella tarbelliana (Grateloup, 1840) (AC coll.).

Cluzaud).

Moulin Debat, Salles, Serravallian, middle Miocene, height $5.0 mm (photo A

THE NAUTILUS, Vol. 120, No. 3

Page 88

B. Landau and C. M. da Silva, 2006

result of the late Neogene cooling events (Monegatti and

Raffi, 2001; Silva and Landau, in press), and does not

survive off West Africa.

Scaphella carlae new species

(Figures 23-29, 49-51)

Description: Shell large, fusiform, of medium thick-

ness, slender and elongate. Protoconch with 1.5 smooth

flattened whorls. Nucleus of medium size, with a small,

blunt-pointed calcarella. Junction with teleoconch not

sharply delimited. Teleoconch with five whorls. The first

teleoconch whorl is short and flat, about three times

wider than tall. Second teleoconch whorl increases in

height rapidly, so that suture, nearly horizontal on first

Ww hel. becomes more oblique. By third teleoconch

whorl, width is 1.5 times height. Abapically the whorls

become taller and more convex, with narrow, slightly

concave sutural ramp. Last whorl about 77% of total

height, elongate, slender and not particularly inflated,

slightly shoul dered in some specimens. Sculpture of very

faint to obsolete spiral threads, most evident on early

whorls and below suture. Aperture 66% of total height,

tall, relatively narrow. Outer lip not thickened, convex in

profile. C olumella weakly concave, bearing 4—5 oblique,

symmetrical, elevated columellar folds of wadiable thick-

ness, increasingly oblique abapically; middle folds most

see dev eloped, abapical fold absent in smaller speci-

mens; folds strongest and somewhat flattened in gerontic

oe Columellar and parietal callus very chin and

worn in most specimens. Siphonal canal long and slightly

recurved abapically. Siphonal fasciole absent.

Type Material: Holotype; IRScNB IST 6994, height,

147.0 mm; paratype; IRScNB IST 6995, height,

147.0 mm.

Type Locality: Velerin conglomerates, Velerin, Este-

pona, province of Malaga, Spain.

Stratum Typicum: Velerin Conglomerates, lower Pi-

acenzian, lower Pliocene.

Other Material Examined: Thirteen specimens,

from the type locality, BLP coll.

Distribution: Lower Pliocene: western Mediterra-

nean, Estepona.

Etymology: Named after Carla Santos, botanist, cur-

rently working at Coimbra University; companion and

moral support to one of the authors (CMS),

Comparative Remarks: —Scaphella carlae new species

differs from the North Sea Basin Pliocene species S.

lamberti in having a more elongate shell, with a higher

spire, the spire whorls are relatively taller and the last

whorl less inflated, the outer lip is convex in profile and

not sinuous as in S. lamberti, and the aperture is rela-

tively smaller and much narrower. The protoconch of S.

carlae consists of 1.5 flattened w hork. with a small blunt

calcarella at the apex. The protoconch of S. lamberti

comprises about two whorls, is smaller (7.3 versus 9.3

mm diameter), rounded, and more bulbous. Marquet

(1997) discussed the variability in height/width ae of S.

lamberti in his Belgian assemblage. Fic ratio was 2.1 to

2.7 (average: 2.4). This is identical to the variability we

have Found (Figure 58) for the specimens of S. lamberti

examined (BLP coll.), 2.1 to 2.6 (average: 2.3). These

contrast markedly with the range observed in S. carlae,

2.7 to 3.1 (average: 2.9).

High-spired and elongate Scaphella bolli from the

middle Miocene North Sea Basin is more similar in

shape to S. carlae than S. lamberti. Indeed, the range of

height/width observed by Marquet (199 7), 2.8 to 3.1 (av-

erage: 2.96) is about the same as that for S. carlae. How-

ever, S. bolli has fewer columellar folds (83—4 versus 4-5),

the spiral sculpture is more strongly developed and the

protoconch is even smaller (5.3 mm diameter), with the

first whorl even more bulbous and the calcarella more

pointed in S. bolli than in S. carlae.

The middle Miocene Atlantic species Scaphella mio-

caenica (Figure 18) from the Loire Basin has a broader,

squatter, more solid shell, with more shouldered whorls.

The protoconch is rather similar in shape to that of S.

carlae, with 1.5 flattened whorls and a small blunt cal-

carella at the apex, but much smaller (7.0 versus 9.3 mm

diameter).

Scaphella tarbelliana (Grateloup, 1840) from the At-

lantic lower and middle Miocene Aquitaine Basin (Fig-

ures 30-31) has characters intermediate between those

of S. carlae and S. lamberti. The protoconch of S. tar-

belliana is about 6.2 mm diameter. It is smaller, narrower

and less inflated than S. lamberti, but still broader, with

a proportionally lower spire than S. carlae. The outer lip

is convex in profile and not sinuous as in S. lamberti, and

Figures 32-51.

Scaphella species. 32-33. Scaphella sp., juvenile (CMS coll.).

Vale de Freixo, Pombal Region, central-western

Portugal. Uppermost Zanclean to lowermost Piacenzian, lower-middle Pliocene, ey 54.6 mm. 34. Scaphe la sp., juvenile (CMS

coll.). Vale de Freixo, Pombal Region, central-western Portugal. Uppermost Zanclean to lowermost Piacenzian, lower-middle

Pliocene, height 31.4 mm. 35-37. Scaphella miocaenica (Fischer and Tournouér, 1579), juvenile (BLP coll.). Ferri¢re-Largon, Loire

Basin, France, Serravallian, middle Miocene, height 19.5 mm. 38-40. Scaphella sp., juvenile (M/V coll.).

Basin, Spain, Zanclean, lower Pliocene, height 34.1 mm. 41-43. Scaphella sp., juvenile (M/V coll.).

Bonares, Guadalquivir

Bonares, Guadalquivir Basin,

Spain, Zanclean, lower Pliocene, height 34.0 mm. 44-45. Scaphella lamberti (Sowerby, 1816), juvenile (BLP coll.). Vrasenedok,

Kallo, Oost-Vlaanderen, Antwerp, Belgium, Oorderen Sands, middle

1816), juvenile (BLP coll.)

mim. 48. Scaphella lamberti (Sowerby, 1816), juvenile (BLP coll.).

Pliocene, height 38. 8 mm. 46-47. Sc aphella ee rti (Sowerby,

_ Vrasenedok, Ki ulo, Oost-Vlaanderen, Antwerp, Be leium, Oorderen Sands, middle Pliocene, height 35.0

Vrasenedok, Kallo, Oost-Vlaanderen, Antwerp, Belgium, Oorderen

Sands, middle Pliocene, height 32.0 mm. 49-51. Scaphella carlae new species juvenile (BLP coll.). Velerin Conglomerates, Velerin,

Estepona, Spain, lower Piacenzian, middle Pliocene, height 47.5 mm.

Page 90

THE NAUTILUS, Vol. 120, No. 3

the aperture intermediate in width between the two.

Nevertheless, the overall outline of S. tarbelliana is still

considerably more inflated than that of S. carlae.

Scaphella miocaenica differs from S. tarbelliana in being

thicker-shelled, squatter, and in having a lower spire and

a more inflated, more shouldered last whorl.

Glibert (1952) noted that even at the juvenile stage S.

miocaenica and S. lamberti were quite different, how-

ever, the character of the juvenile shell was not consid-

ered by subsequent authors. Certainly the juvenile shell

of S. miocaenica is quite different from that of S. lam-

berti and S. carlae, being much broader, and the proto-

conch whorls more depre ssed. The shape of the juvenile

shell of S. carlae is similar to S. lamberti, but in the latter

the apex is more rounded, the calcarella more elevated

and the diameter of the first teleoconch whorl is smaller.

Scaphella sp.

(Figures 38-43, 52-57)

Description: Shell large, fusiform, relatively thick-

shelled. Protoconch with about 1.5 smooth, flattened

whorls, with small, blunt-pointed calcarella. Junction

with teleoconch not sharply delimited. Teleoconch with

3-3.5 whorls. First eae whorl short, flat, width

about three times height. Second teleoconch whorl

weakly angular in profile, increasing in height rapidly, so

that suture becomes more oblique than on first whorl.

Last whorl about 86% of total height, inflated, shoul-

dered weakly a short distance below suture. Sculpture of

very faint to obsolete spiral threads, most evident on

early whorls and below suture. Aperture 74% of total

height. Outer lip broken in examined specimen. Col-

umella almost straight, bearing four narrow, elevated,

oblique columellar folds, increasingly oblique abapically:

adapical three folds of roughly equal strength, abapical

fold much weaker. Columellar and parietal callus not

preserved. Siphonal canal long and straight. Siphonal fas-

ciole absent.

Dimensions and Material: Maximum height: 112.0

mm (although incomplete specimens suggest up to ap-

proximately 130 mm). Four specimens, BLP coll.; six

specimens, M-V coll. All from Bonares, Guadalquivir Ba-

sin, Huelva, Spain. Zanclean, lower Pliocene.

Discussion: Despite intensive collecting by one of the

authors (BL) and dedicated local collectors Manuel Mo-

lin and Daniela Velo for more than 30 years, only six

incomplete adults and two juveniles have been found in

the ¢ Guadalquivir Basin de »posits. However, it is clearly

not conspecific with Scaphella carlae. The lower

Pliocene Atlantic specimens from the Guadalquivir Basin

are thicker-shelled (maximum shell thickness 4.9 mm),

squatter, with fewer whorls, the last whorl distinctly

shouldered as opposed to convex or weakly shoulde red,

and there are only four columellar folds, which are nar-

rower than in the coeval Mediterranean S. carlae. The

maximum diameter of the protoconch (9.7 mm) is similar

to that of S. carlae.

Of all the European species of Scaphella, the Guad-

alquivir Basin shells are most similar to the Atlantic

middle Miocene Serravallian S$. miocaenica from the

Loire Basin of France. This latter species is also relatively

squat, thick-shelled, with the last whorl shouldered, and

with four folds on the columella. The protoconch diam-

eter of Scaphella sp. is, however, aoe than in S. mio-

The juvenile stage of S.

miocaenica is quite ohafdoterisae: the last whorl very

broad (Figures 35-37), unlike that of S. lamberti or S.

caenica (9.7 versus 7.0 mm)

‘igures 52-57. Scaphella sp. (BLP coll.), Bonares, Guadalquivir Basin, Spain, Zanclean, lower Pliocene, 52-54. Height 98.7 mm.

55 56. Height 112.2 mm. 57. Height 90.4 mm

B. Landau and C. M. da Silva, 2006

Page QI

carlae. Fortunately, two perfectly preserved juvenile

specimens are av ailable to us collected by Manuel Molin

and Daniela Velo. The shape of the juvenile shell is most

similar to that of S. miocaenica, possibly even more in-

flated, but larger at the same number of whorls, and the

diameter of the protoconch and first teleoconch whorls is

greater.

The specimens from the lower Pliocene Mondego Ba-

sin of Portugal (Silva, 2001) are both juvenile (Figures

32-34). Their protoconch characteristics and ae shape

of the juvenile shell are more similar to those of

Scaphella sp. than S. carlae, and they have provisionally

been added to the distribution.

CONCLUSIONS

In the Recent volutid faunas, non-planktotrophic larval

development most likely induces the establishment of

locally distinct populations, which ultimately results in

poorly defined species-group taxa, as with the Cymbiola

Total width / total height

80 =

70 -

Total width

|_|

110 120 130 140 150 160

Total height

Apertural height / total height

90 -

Apertutal height

80 +

110 120 130 140 150 160

Total height

4 Scaphella lamberti ® Scaphella carlae

Figure 58. Morphometric comparisons between Scapliella

lamberti and S. carlae new species. Measurements in mm

pulchra group from Queensland, Australia (Darragh and

Ponder, 1998).

The same applies to Recent Scaphella species in the

New World, which explains the enormous ia »pancy in

the number of species recognized by different authors

(four, Weaver and du Pont, 1970: 12, Clench, 1946). A

similar situation occurs with European Scaphella, the

species being restricted both geographically and _strati-

graphically. This is illustrate .d by the presence of two

different species in the Atlantic Tee Loire and

Aquitaine Basins and the description of two coeval, but

distinct species present on either side of the Straits of

Gibraltar in the Pliocene. Most of the European species

are relatively easily characterized, although the middle

Miocene to lower Pliocene (lower to upper Redonian)

Scaphella species from the Atlantic Loire Basin, north-

ern France are less clearly distinguished. Brébion (1964)

recorded the presence of both S. miocaenica and S. lam-

berti in these deposits and observed a transition from one

to the other from the middle Miocene to the Pliocene.

This study of European Scaphella revealed that the

genus as traditionally regarded is heterogeneous, with

le sar and consistent differences in shell morphology be-

tween Old and New World species. Old World Scaphella

occur continuously in the European Cenozoic from the

Paleocene to the middle Pliocene and form a distinct

group from the Neogene to Recent New World species.

Assuining that Bondarev (1997) and Bandel (2003) are

correct, and that that the Scaphellinae have a Cretaceous

Tethyan origin, and that Dall (1907) is also correct, and

that ‘Scaphella originated from Caricella, it is hypoth-

esized that the New World Paleocene Sc aphella originat-

ed from a Cretaceous/Paleocene Old World ancestor,

which emigrated to the New World from the Tethys, and

that the Neogene New World Scaphella species are a

distinct stock, which evolved from Caricella. This would

explain the similarities between Old and New World Pa-

leocene Scaphella and the differences with the New

World Neogene stock. The European Paleocene to

Pliocene species traditionally assigned to Scaphella

therefore would constitute a separate stock from those in

the New World, and might even correspond to a distinct

genus-level taxon.

ACKNOWLEDGMENTS

We would like to thank Dr. Alan Beu of the Institute of

Geological and Nuclear Sciences (New Zealand) and

Geerat Vermeij of the Dep: aoe nt of Geology, Univer-

sity of California at Davis (USA) for their critical review

of the manuscript. Chris Garvie of the Texas Memorial

Museum, Texas (USA) for advice and photographs of the

Paleocene North American Scaphellinae. Phils Diegel

of Florida (USA) for supplying specimens ‘of juvenile

American fossil S¢ aphu lla. Manuel Molin and Daniela

Velo of Bonares (Spain), Robert Marquet (Belgium) and

Alain Cluzaud (France iollawe dus access to the! ir collec-

tions and preven specimens for this work. Inge-

nae 09:

Page 92

THE NAUTILUS, Vol. 120, No. 3

mann Schnetler of Langa (Denmark) for photographs of

Danish Paleocene specimens. Contribution of the Por-

tuguese FCT Project POCTI 32724/99—Comparative

(palaco)environmental analysis of oceanic and coastal do-

mains, over the last 20 Ma, based on calcareous nanno-

plankton (CANAL), co-financed by European Union's

FEDER.

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THE NAUTILUS 120(3):94—100, 2006

Page 94

A new record of introduced Cerion (Gastropoda: Pulmonata:

Cerionidae) in southeastern Florida

M. G. Harasewych

Department of Invertebrate Zoology

National Museum of Natural History

Smithsonian Institution

Washington, DC 20013-7'

[email protected]

Josiah Strauss

012 USA

Department of Geology and Geophysics

Texas A&M University

College Station, TX 77843-3115 USA

[email protected]

ABSTRACT

A large, well-established colony of the land snail genus Cerion

is reported from Delray Beach, Florida, far north from the

range of the only native species in Florida, Cerion incanum

(Binney, 1851) and its subspecies, and from the well-

een populations introduced by Paul Bartsch during

the early 20" Century. Although hurricanes have frequently

been proposed as agents of long distance dispersal of Cerion

species, the Delray Beach C solony is more probably the result of

intentional introductions during the mid-20'" C ventury. The

morphology of the shells in this colony is uniform except for the

degree of pigmentation, yet it does not match precisely any of

the fourteen named taxa from the Little Bahama Bank or the

Bimini Islands, the nearest and most probable sources of the

introduced propagules. Rather, the phenotype appears to com-

bine elements of several taxa from both island groups, suggest-

ing that the Delray Beach Colony may be a result of hybrid-

ization among animals from multiple and disparate introduc-

tions.

Additional Keywords: Introduced species, land snails, hybrid-

ization, Bahamas.

INTRODUCTION

The land snail genus Cerion has long been known for its

extreme morphologic: al dive orsity, especli ally in the faunas

of Cuba, the Bahamas, and Cayman Islands (see Wood-

ruff, 1978, and references therein). Although the oldest

Cenozoic records of the genus are from the Oligocene of

western Florida (Table 1A; Figure 1, W), Cerion is lim-

ited to a single species, Cerion incanum (with four sub-

species or forms, see Table 1B; Figure 1, ®) in the

Recent fauna of Florida (Pilsbry, 1946: 163, fig. 77). The

species inhabits nearshore vegetation from Key Biscayne

southward throughout the Florida Keys to Key West.

Several authors (e.g., Binney, 1851: 153; Pilsbry, 1902:

213; 1907: 193; 1946; 162; Dall, 1905a: 30) have sug-

gested that C. incanum is not directly descended from a

Florida Oligocene species but rather from Cerion that

re-colonized the Florida Keys from Cuba following the

interglacial high stands that had completely submerged

southern Florida during the Pleistocene (Hearty et al.,

1999). Pilsbry (1902: 213) regarded Cerion incanum to

be most closely related to the Cuban C. maritinum

(Pfeiffer, 1839).

The Cerion fauna of Florida had been significantly

increased through a series of well-documented experi-

mental transplantations of non-native species begun by

Bartsch in 1912 (see Bartsch, 1912; 1920; 1949, ane ref-

erences therein). Bartsch’s initial introductions were

contined to two species that he subsequently named Ce-

rion casablancae Bartsch, 1920, and Cerion viaregis

Bartsch, 1920, both from Andros Island, Bahamas. Each

of these was introduced to several of the Florida Keys,

including the Dry Tortugas that were thought to be de-

(Table 1C). Bartsch later ex-

panded his experiments to include introductions of an

additional dozen species from Cuba, Puerto Rico,

Curacao, and several Bahamian islands, to sites in the

Dry Tortugas, especially Fort Jefferson on Garden Key,

and Loggerhead Key. He monitored and reported on

these introduced colonies for decades. Most of the in-

troduced colonies have since died out, but several (e.g.,

Cerion casablancae on Indian Key) flourish to this day,

and some (e.g., C. viaregis on Newfound Harbor Key

and C. casablancae on Bahia Honda Key) have hybrid-

ized with the native C. incanum (see Bartsch, 1920:

Woodrutf and Gould, 1987)

We have recently encountered a sizeable colony of

Cerion in Delray Beach, Florida (Figure 1, *), far north

of the known distribution of native Cerion incanum (Fig-

ure |, ®) and the sites of Bartsch’s introductions (Figure

1, ®). The Delray Beach Colony inhabits vegetation on

seaside sand dunes between the Atlantic Ocean and

Route ALA, from Atlantic Avenue (26°27.696' N,

$0°03.483' W) northward to near the northern limit of

the Gulfstream Golf Course (26°29.284' N, 80°03.218'

W), where the dunes and vegetation are replaced by

private homes that extend to the beach. These Cerion are

most abundant just south of George Bush Boulevard

(26°28.350' N, 80°03.373' W), and near the northern end

void of Cerion incanum

M. G. Harasewych and J. Strauss, 2006

Page 95

Table 1. The family Cerionidae in Florida.

A. Fossil Taxa (all Oligocene, from Balast Point, 1

Cerion (Eostrophia) anodonta Dall, 1890.

Cerion (Eostrophia) anodonta var. floridanum Dall, 1890.

Microcerion floridanum Dall, 1915.

B. Living, Native Taxa (with their type localities).

Cerion incanum incanum (Binney, 1851). Key West.

Cerion incanum fasciatum (Binney, 1859). Key Biscayne.

Cerion incanum saccharimeta ‘Blanes’ Pilsbry ‘and Vanatta,

Cerion incanum vaccinum Pilsbry, 1902. Key Vacca.

‘ampa Bay)

1899. Sugarloaf Key.

C. Non-native species of Cerion introduced intentionally to the Florida Keys and the Dry Tortugas by Bartsch, 1912-1924.

Cerion viaregis Bartsch, 1920. From Andros, Bahamas, introduced to: Second Ragged Key North of Sands Key; Tea

Table Key; Duck Key; Newfound Harbor Key; Key West; Boca Grand Key: € Garden Key; Loggerhead Ke Y; Man Key;

Boy Key.

Cerion casablancae Bartsch, 1920, From Andros, Bahamas, introduced to: First Ragged Key North of Sands Key;

Sands Key; Indian Key; Bahia Honda Key;

Loggerhead Key.

Cerion (P aracerion) tridentatum Pilsbry and Vanatta, 1895. From Cuba introduced to: Key West; Garden Key, north

side of the parapet at Fort Jefferson; Loggerhead Key.

Cerion mumia (Sowerby, 1834). From the point at Miramar, Cuba, introduced to: Garden Key, north side of the para-

pet at Fort Jefferson, Loggerhead Key.

Cerion chrysalis (“Ferussac” Beck, 1837). From near Cabanas F

the parapet at Fort Jefferson, Loggerhead Key.

Fort, Cuba, introduced to: Garden Key, north side of

Cerion sculptum (Poey, 1858). From near the lighthouse at Mariel, Cuba, introduced to; Garden Key, north side of the

parapet at Fort Je fferson.

Cerion “n. sp.” Young. From East of the point at Mariel, Cuba, introduced to: Garden Key, north side of the parapet

at Fort Jefferson.

Cerion sp. {small, mottled]. From Andros, Bahamas, introduced to: Bird Key.

Cerion sp. Mottled. From New Providence, Bahamas, introduced to: Logge rhead Key,

Cerion crassilabris (“Shuttleworth” Sowerby, 1875). From Puerto Rico, introduced to: Loggerhead Key.

Cerion uva (Linne, 1758). From Curagao, introduced to:

Loggerhead Key.

Cerion “mayori” (not clear if this is a typographical error for C. mayoi Maynard and Clapp, 1920, or a manuscript

name). From Middle Bight, Andros, introduced to: Loggerhead Key.

Unspecified Cerion. From Puerto Rico and “El Salvador,”

introduced to: Loggerhead Key:

Cerion incanum (Binney, 1951). Source unspecified [Florida Keys], introduced to: Logge head Key, Man key, Boy Key.

of Gulfstream Golf Course (26°29.247' N, 80°03.229'

W). where densities exceeded 15 individuals per square

meter. The snails in this colony are fairly uniform in their

shell morphology (Figures 3, 5-8), but vary in degree of

pigmentation from pure white (Figure 7) to wealcly { Fig-

ure 6, S) or strongly (figures 2, 4) mottled with oblique

bands of brown.

Many Cerion researchers (e.g., Pilsbry, 1907; Mayr

and Rosen, 1956; Clench, 1957; Gould and Woodruff,

1978) accepted the hypothesis that hurricanes play a sig-

nificant role in the long distance dispersal of Cerion spe-

cies and are a major factor in determining biogeographic

patterns within the genus. A review of iistomes! hurri-

cane tracks [http://hurricane csc. noaa.gov/hurricanes/

viewer.html| revealed that the Little Bahama Bank and

the Bimini Islands (both within 120 km of Delray Beach)

are the closest potential sources for this colony under this

dispersal scenario. However, intentional human intro-

duction during the mid- 20th Century is, IN our view, a

more credible explanation for the origin of this colony.

Dr. Edward Petuch (personal communication) re-

counted to us his conversations with the late P. L. and

T. L. McGinty, in which they acknowledged introducing

at least six different Bahamian populations of Cerion,

“mostly from West End (Grand Bahama Island) and the

Bimini Islands,” to the vicinity of their home in Boynton

Beach. These second-hand accounts of Cerion introdue-

tions are given credence by an earlier report of an

anomalous “population of another land snail, Liguus fas-

ciatus (Miiller, 1774), from nearby Boynton Beach

(Craig, 1973) that was subsequently documented as be-

ing ie descendents of an intentional introduction by the

McGinty brothers (Krieger and Austin, 1975). W yhether

transported by a hueeioane or intentionally introduced,

the most probable sources for the Delray Beach colony

are the Cerion faunas of the Little Bahama Bank or of

the Bimini Islands.

Nine species or subspecies of Cerion have been de-

scribed from the Little Bahama Bank (Clench, 1957)

(Table 2, Figures 9-18). Gould and Woodruff (1975)

reduced these taxa to two semispecies, C. abacoensis

Pilsbry and Vanatta, 1895, and C. bendalli Pilsbry and

Vanatta, 1896, based on morphometric and allozyme

studies, but noted that the various shell morphotypes

could be localized to specific regions on the Little Ba-

hama Bank.

Clench (1935; 50) conjectured that Cerion milleri

(Pfeiffer, 1867), described from Duck Key, Exuma

Page 96

THE NAUTILUS, Vol. 120, No. 3

Figure 1.

Cerion incanum and subspecies (diamonds,

Distribution of the Genus Cerion in Florida. Late Oligocene-Early Miocene species (triangle, V

@). Non-native species of Cerion intentionally introduced to io Florida Keys and the

& Bimini —

Islands

). Recent, native species,

Dry Tortugas by Bartsch, 1912-1924 (circles, @). New record of introduced Cerion (cross, #). Probable sources of the introduction

are the Little Bahama Bank and/or the Bimini Islands. $ Satellite image modified from that av ailable at http://visibleearth nasa.gov.

Group, might actually be from Duck Key in Cherokee

Sound, off Abaco, noting that he could not find any

“Duck Key” in the Exuma Group on modern maps.

Gould and Woodruff (1978: 381) commented that if

Clench were correct, C. milleri would become the senior

synonym of C. bendalli, but noted that they could not

verify Pfeiffer’s locality, and that similar mottled Cerion

occur throughout the Bahamas, inc luding the Exumas.

Perusal of Nautical Chart 26-D ( (Department of the Navy

Hydrographic Office, 1965) reveals Duck Cay to be lo-

cated west of the southem end of Great Exuma Island

(23°27.22' N, 76°02.60' W), supporting Pfeiffer’s origi-

nal type locality and excluding this taxon from the fauna

of the Little Bahama Bank.

The Cerion fauna of the Bimini Islands consists of five

named taxa (Table 3, Figures 19-23). Clench (1942) had

synonymized three of the taxa (Cerion pillsburyi Pilsbry

and Vanatta, 1897, C. canonicum Dall, 1905, and C.

northropi Dall, 1905), but later (Clench, 1956) he named

another taxon. Mayr and Rosen (1956) concluded that it

was not possible to classify the existing colonies simply in

terms of three nominal species, and noted that some

colonies formed highly variable hybrid populations.

The transplantation experiments of Bartsch (1920)

documented that colonies resulting from single introduc-

tions retain the morphology of heir parental stock for

multiple gene rations following transplantation. However,

most Cerion “species ” hybridize freely when com-

mingled. In the short term, such hybrid populations pro-

duce morphological and mole cular features not evident

in either parent population (Bartsch, 1920; Woodruff,

1989). Over a period of decades, the hybrid phenotype

and genotype may be gradually assimilated into that of

the numerically dominant taxon (Woodruff and Gould,

1987), yet even after millennia, traces of past hybridiza-

tion may persist as area effects (C Goodfriend and Gould,

1996).

Shells from the Delray Beach colony do not resemble

precisely any of the fourteen named taxa from either the

Bimini Islands or the Little Bahama Bank as might be

expected if this colony were the result of a single intro-

duction. Rather, De lray Beach colony Cerion e hibit at-

M. G. Harasewych and J. Strauss, 2006

Figures 2-8. Five specimens of introduced Cerion from northern end of Delray Park, Delray Beach, Florida (26°28.23' N,

§0°03.39' W), USNM 1086626. 2. Specimen A. 3. Specimen A coated with ammonium chloride to show sculptural details. 4.

Specimen B. 5. Specimen B coated with ammonium chloride to show sculptural details. 6-8. Specimens C-E, weakly pigmented or

unpigmented specimens

Figures 9-15. ’ 337

Abaco, Bahamas. 10. Cerion maynardi Pilsbry and Vanatta, 1895, lectotype, ANSP 25338, Abaco, Bahamas. 11-12. Cerion aba-

coensis bendalli Pilsbry and Vanatta, 1896, 11. Lectotype, ANSP 25343, Great Abaco, Bahamas. 12. Lectotype cv vated with ammo-

nium chloride to show sculptural details. 13. Cerion oweni oweni Dall, 1905, lectotype, USNM 179436, Little Abaco, opposite Marsh

Harbor and Riding Point, Grand Bahama. 14. Cerion oweni reticulatum Dall, 1905, lectotype, USNM 179443, Sugar Loaves, Abaco

15. Cerion oweni incisum Dall, 1905, lectotype, USNM 179440, Stranger Cay, NW of Little Abaco. 16. Cerion oweni vermiculim

Dall, 1905, lectotype, USNM 179442, Mathews Point, south side of Great Abaco Bahamas. 17. Cerion chrysaloides Plate, 1907, MCZ

116008, Eight Mile Rock, Grand Bahama Island. 18. Cerion lucayanorum Clench, 1938, paratype, MCZ 116015. NW portion of

Mores Island, 32 miles NW of Southwest Point, Great Abaco Island, Bahama Islands. Scale bar = 1 cm for all specimens

Cerion taxa from the Little Bahama Bank. 9. Cerion abacoensis Pilsbry and Vanatta, 1$95, lectotype, ANSP 25337

Page 95

THE NAUTILUS, Vol. 120, No. 3

Table 2. Cerion taxa described from the Little Bahama Bank, listed chronologically, together with their type localities, primary type

specimens, and subsequent notes on distribution,

Cerion abacoensis Pilsbry and Vanatta, 1895. Abaco, Bahamas. Lectoty ie ANSP 25337a (Baker, 1963: 206), now ANSP 25337,

4 paralectotypes ANSP 411926. Gould and Woodruff (1978: 3

ing, Great Abaco.

79, fig. | report that this phenotype is restricted to the Cross-

Cerion maynardi Pilsbry and Vanatta, 1895, Abaco, Bahamas. Lectotype, ANSP 25335a (Baker, 1963: 206), now ANSP 25338, 4

paralectotypes ANSP 411925. Gould and Woodruff (1978: 379) reported finding this phenotype only near Hole-in-the-Wall

Light, on Great Abaco.

Cerion abacoensis bendalli Pilsbry and Vanatta, 1896. Great Abaco, Bahamas. Le ctotype, ANSP 25343a (Baker, 1963: 206), now

ANSP 25343. Gould and Woodruff (1978: figs. 1-3) report that this phenotype is broadly distributed on Grand Bahama Is-

land, Little Abaco, and northern and central Great Abaco.

Cerion oweni oweni Dall, 1905. South side of Little Abaco, Bahamas. Lectotype,

USNM 179436 [originally illustrated specimen

(Dall, 1905b: pl. 58, fig. 12) is here designated as the lectotype]|, 17 paralectotypes, USNM 1086731. Dall (1905b: 443) re-

ported the typical form to occur on Little Abaco, Great Abaco, and Grand Bahama Island. The type locality is that of the

lectotype.

Cerion oweni reticulatum Dall, 1905. Sugar Loaves Rocks, NW of Elbow Cay, Great Abaco, Bahamas. Lectotype, USNM

179443 [originally illustrated specimen (Dall, 1905b: pl. 58, fig. 8) is here designated as the lectotype], 13

USNM_ 1086732.

Cerion oweni incisum Dall, 1905. Stranger Cay Beach, N

paralectotypes,

NW of Little Abaco. pune (ie USNM 179440 [originally illustrated

specimen (Dall, 1905b: pl. 55, fig. 10) is here design: ii ‘das the lectotype},

paralectotypes, USNM 1086733.

Cerion oweni vermiculum Dall, 1905, Mathews Point, south side of Great roe Bahamas. Lectotype USNM 179442 ae

illustrated specimen (Dall, 1905b: pl. 58, fig. 3) is here designated as the lectotype], 3 paralectotypes, USNM 10867:

Cerion chrysaloides Plate, 1907. Eight Mile Rock, Grand Bahama Island, MCZ 116008,

Cerion lucayanorum Clench, 1938. NW portion of Mores Island, :

Islands, paratype, MCZ 116015.

2 miles NW of Southwest Point, Great Abaco Island, Bahama

tributes of several of the potential source populations.

The broad shells with fewer, taller whorls, a longer, more

acute conical portion of the spire, and thin parietal callus

are most similar to C. pillsburyi and C. eximium lerneri

from the Bimini Islands. However, the strong ribbing,

patterns of pigmentation, as well as the shape, size, and

orientation of the aperture and columella more closely

resemble taxa from the Little Bahama Bank, particul: uly

Figures 19-23. Cerio taxa from the Bimini Islands, Bahamas. 19. Cerion pillsburyi Pilsbry and Vanatta, 1897, lectotype, ANSP

72136a. Gun C:

20. Cerion canonicum Dall, 1905, lectotype, USNM 127460. Gun Key, 21. Cerion northropi Dall, 1905, lectotype,

USNM 125135 sack of the westernmost ni ts near Gun Cay. 22. Cerion biminicnse Henderson and Clapp, 1913, lectotype, USNM

252849. Southern end of North Bimini C

23. Cerion eximium lerneri Clench, 1956, holotype, MCZ 186530. East Bimini

M. G. Harasewych and J. Strauss, 2006

Page 99

Table 3. Cerion taxa described from the Bimini Islands, listed chronologically, together with their type localities, primary type

specimens, and subsequent notes on distribution.

Cerion pillsburyi Pilsbry and Vanatta, 1897. Gun Cay,

Bahamas. Lectotype, ANSP 72136a (Baker, 1963: 207), now

ANSP 72136, 3 paralectotypes ANSP 411921.

Cerion canonicum Dall, 1905. Gun Cay, Bahamas. Lectotype, USNM 127460 [originally illustrated specimen (Dall, 1905b: pl.

5S, fig. 13) is here designated as the lectotype]|, 4 paralectotypes, USNM 1086735.

Cerion northropi Dall, 1905. Bahamas, probably one of the westernmost islets near Gun Cay. Hestoyye, USNM 12413 oo

nally illustrated specimen (Dall, 1905b: pl. 5S, fig. 11) is here designated as the le sctotype], 2

Cerion biminiense Henderson and Clapp, 1913. Southern end of North Bimini Cay, Bahamas. Lectotype, t

2 paralectotypes USNM 10867.

ISNM 252849 en

nally illustrated specimen (Henderson and Clapp: 1913, pl. 4 fig. 10) is here designated as the lectotype], 158 paralectotypes,

USNM 1086737.

Cerion eximium lerneri Clench, 1956. Holotype, MCZ 186830. Southern tip of East Bimini, Bimini Islands,

Bahamas.

C. abacoensis and C. oweni oweni. Shell morphology

suggests that the Delray Beach colony may be the result

of ‘hybridization of snails from two or more populations

introduced from the northern Bahamas some time near

the middle of the twentieth century. Future genetic

analyses may aid in the identification of the source Ba-

hamian popul: ition or populations for the Delray Beach

colony.

ACKNOWLEDGMENTS

We are most grateful to Dr. Edward J. Petuch of Florida

Atlantic University for sharing his recollections and rem-

iniscences of conversations with the McGinty brothers,

and for taking part in initial surveys for introduced popu-

lations of Cerion between Manalapan and southern Del-

ray Beach, Florida. We also thank Paul Callomon, De-

partment of Malacology, Academy of Natural Sciences of

Philadephia, and Adam Baldinger, Department of Mala-

cology, Museum of Comparativ e Zoology, for the loan of

type specimens in their care. This re »search was sup-

ported by NSF Grant # EAR 0106936.

This is Smithsonian Marine Station at Fort Pierce

Contribution Number 653.

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Pilsbry, H. A. 1946. Land Mollusca of North America. The

Academy of Natural Sciences of Philadelphia, Monograph

3, Volume 2 , part 1: viii + 520 pp.

Woodruff, D. S. 1978. Evolution and adaptive radiation of Ce-

rion: a remarkably diverse group of West Indian land

snails. M: alacologia 17: 223-239.

Woodruff D. S. 1989. Genetic anomalies associated with Ce-

rion hybrid zones: the origin and maintenance of new

electromorphic variants called hybrizymes. Biological

Journal of the Linnean Society 36: 281-294.

Woodruff, D. S. and S. J. Gould. 1987. Fifty years of interspe-

cific hybridization: genetics and morphometrics of a con-

trolled experiment on the land snail Cerion in the Florida

Keys. Evolution 41: 1022-1045,

THE NAUTILUS 120(3):101—105, 2006

Page 10]

A new early Miocene Muracypraea Woodring, 1957

(Gastropoda: Cypraeidae) from the Pisco Basin of southern Peru

Thomas J. DeV ries!

Burke Museum of Natural History

and Culture

University of Washington

Seattle, WA 9S195 USA

[email protected]

Lindsey T. Groves

Malacology Section

Natural History Museum of

Los Angeles County

900 Exposition Boulevard

Los Angeles, CA 90007 USA

Mario Urbina

Museo de Historia Natural

Universidad Nacional Mayor de San

Marcos

Departamento de Paleontologia de

Vertebrados,

Avenida Arenales 1256, Jésus Maria,

Apartado 14-0434

Lima 14, PERU

ABSTRACT

The cypraeid gastropod, Muracypraea ormenoi new species, is

described from lower to middle Miocene beds of the upper-

most Oligocene to middle Miocene Chilcatay Formation, Pisco

Basin, southern Peru. It is the first cypraeid reported from

southern Peru and represents the southernmost occurrence of

this genus. Muracypraea is normally a warm-water gastropod

and its occurrence in the study area indicates dispersal by

means of coastal countercurrents into waters possibly cooled by

coastal upwelling. The new species shares a neritic habitat with

endemic molluscan taxa and with taxa ranging northward from

the Navidad Basin of central Chile.

INTRODUCTION

Muracypraea ormenoi new species is described from the

Chilcatay Formation of the Pisco Basin, southern Peru.

Associated mollusks and microfossils from correlative

nearby strata indicate an age of approximately 20 Ma

(early Miocene) for its occurrence. The new species is

represente -d by a well preserved holotype and a poorly

preserved specimen. This new species represents the

southernmost occurrence of Murac ypraea Woodring,

1957, and indicates a dispersal into waters of eiewal:

coastal upwelling environments. The new species also

shares a neritic habitat with endemic molluscan taxa and

taxa ranging northward from the Navidad Basin of cen-

tral C fle (D DeVries and Frassinetti, 2003). Its occurrence

at 15°S also accentuates the modern-day diminishment

of the range of Muracypraea since the early and middle

Miocene, because M. mus (Linnaeus, 1758), the last liv-

ing representative of the genus, is restricted to the south-

western Caribbean coasts of Venezuela and Colombia

(Lorenz and Hubert, 2000).

' Mailing address: Box 13061, Burton, WA 98013 USA

STRATIGRAPHY AND AGE

The Pisco Basin is a foreare basin that extends along the

narrow coastal plain of southern Peru from Paracas to

Nazca, Ica De citags nt (Figure 1). Figure 2 indicates the

type locality of the new species. Cenozoic marine beds

are exposed throughout the coastal desert, including

those of the upper Oligocene to middle Miocene Chil:

catay Formation and the overlying lower middle Mio-

cene through middle upper Pliocene Pisco Formation

(DeVries, 1998).

CHILCATAY FORMATION

The name Chilcatay Formation was assigned to upper

Oligocene to lower Miocene strata near Pampa Chil-

catay, approximately 60 km west of Ica, by Dunbar et al.

(1990). The Chilcatay Formation consists of basal sand-

stones associated with a transgression at approximately

25 Ma (DeVries, 2001), tuffaceous and diatomaceous

siltstone indicative of shelf depths and a coastal-up-

welling regime (Dunbar et al., 1990), and intercalated

coarse- grained : sandstone that may represent short-lived

early Miocene eustatic sea-level events (DeVries, 1998).

Macharé and Fourtanier (1987) estimated the. strati-

graphic thickness of the C hilcatay Formation at approxi-

mately 250 m.

Pisco FORMATION

Unconformably overlying the Chilcatay Formation is the

Pisco Formation of Adams (1909), named for steeply

dipping, white and yellowish rocks exposed at the end of

the Huamani bridge over the Rio Pisco north of Pisco,

Peru. Dunbar et al. (1990) re ported that the age of the

Formation is late middle Miocene through Pliocene (4—

12 Ma), based on K/Ar dates combined with siliceous

microfossil zonation, whereas DeVries and Schrader

(1997), DeVries (1998), and DeVries and Frassinetti

(2003) assigned an early middle Miocene to middle late

Page 102

THE NAUTILUS, Vol. 120, No. 3

75°W

AMERICAN

HIGHWAY

Paracas

Peninsula

14°S

LOCATION

OF FIGURE 2

Pisco Basin

0 100 km

= =

SCALE

EOCENE OLIGOCENE MIOCENE

Figure 1. Index map showing location of the Pisco Basin in southern Peru and chronostratigraphy of the Pisco Basin (DeVries and

Schrader, 1997).

Pliocene age to the bioclastic conglomerates, tuffaceous

siltstones, and diatomaceous siltstones based on diatom

biostratigraphy and their revised molluscan biostratigra-

phy. The formation varies in thickness throughout the

Pisco Basin from about 200 to 1000 m thick (Dunbar et

al., 1990).

Abbreviations: Abbreviations used for institutional

catalog and/or locality numbers are as follows: DV, col-

lecting localities of Thomas J. DeVries; LACMIP, Natu-

ral History Museum of Los Angeles County, Invertebrate

Paleontology Section; USNM, National Museum of

Natural History, Smithsonian Institution, Washington, DC.

Measurement parameters are defined as follows:

length = greatest distance between anterior and posterior

termini, width = greatest distance between lateral mar-

gins, and height = greatest distance between base and

dorsum. The systematic classification herein follows that

of Schilder and Schilder (1971) with modifications by

Kay (1996).

SYSTEMATIC PALEONTOLOGY

Superfamily Cypraeoidea Rafinesque, 1815

Family Cypraeidae Rafinesque, 1815

Subfamily Bernayinae Schilder, 1927

Genus Muracypraeca Woodring, 1957

Type Species: = Cypraca mus Linnaeus, 1755, by origi-

nal designation. Recent, Venezuela and Colombia.

Diagnosis: Medium to large size, pyriform or triangu-

lar shaped posterior portion of dorsal surface smooth,

warty, or bituberculate, or rarely a central “spike-like”

dorsal tubercule present; labial lip wide, slightly con-

stricted near anterior end, teeth moderately strong; col-

umella wide, teeth strong to weak or absent; aperture

wide, curving in posterior direction toward columella;

fossula indistinct, wide, shallow, smooth; anterior termi-

nal ridges strong and may extend forward in a flattened

or flange-like manner; terminal canals prominent.

Geologic Range:

cent.

Early Miocene (Aquitanian) to Re-

)Cerro

pia Bruja

/ \

K ¢ . f o

{ = RC l, )

\ SY ADV 353-1) LACMIPS

a“ ‘ie, C J e 17783

Yesera Wy e.

SAM

de Amara‘\)

( ae c~)\ Cerros\

\ “Los Tres

. \ Pyramides

Oo ° C

ee % al

SCALE v 7

2 s

Contour interval is LOO m. id

Figure 2. Index map showing the type locality of Mura-

cypraca ormenoi new species at Ullujalla West (LACMIP loc.

17783), and locality DV 553-1 (LACMIP loc. 17615), Dashed

lines indicate roads.

T. DeVries et al., 2006

Page 103

Geographic Range: Miocene: Dominican Republic,

Haiti, Trinidad, Cuba, Jamaica, Venezuela, Colombia,

Panama, Brazil, Ecuador, Peru, Costa Rica, Michoacan,

Mexico, and Baja California Sur, Mexico; Miocene/

Pliocene: Imperial County, California, USA; Pliocene:

Venezuela, Panama, Ecuador; Pleistocene: Venezuela:

Recent: Venezuela and Colombia.

Remarks: In a monograph of the Miocene Gatun For-

mation of Panama, Woodring (1959) acknowledged the

variability of Muracypraea henekeni (Sowerby, 1850) and

the excess of names applied to this species. He stated

that “some of the names in the synonymy may prove to

be usetul for local populations when ade quate samples

are available.” Groves (1997) reviewed the cypraeiform

gastropods of northwestern Ecuador and discussed the

stratigraphic and geographic distribution of the M.

henekeni group in Panama, Ecuador, and Colombia.

Groves (1997, 1998) proposed that a distinctly rhomboi-

dal, bitube rculate lineage be recognized in the western

Caribbean and tropical eastern Pacific that included

specimens previously assigned to M. henekeni (Marks,

1951; Olsson, 1964), as we I as the Pliocene Ecuadorian

species, M. cayapa (Pilsbry and Olsson, 1941).

Muracypraea ormenoi new species

(Figures 3-5)

Diagnosis: A medium-sized Muracypraca of weakly

rhomboidal shape; outer lip extended posteriorly well

beyond inner lip; dorsum bituberculate with posterome-

dial depression.

Description: Shell medium in size; slightly rhomboi-

dal shape; dorsum moderately arched, maximum height

(apex) slightly posterior to midpoint; base slightly flat-

tened; surface generally smooth; lateral margin of right

Figures 3-5.

5. Labral lateral view.

Muracypraea ormenoi new species, holotype LACMIP no. $197, length = 64.9 mm. 3. Dorsal view. 4. Apertural view

Page 104

THE NAUTILUS, Vol. 120, No. 3

side weakly corrugated ventrally; dorsum with broad, low

tubercles: tubercle on labral side situated axially between

apex and periphery; tubercle on columellar side smaller

and situated slightly more posteriorly and closer to axial

median: medial tubercle at apex absent or broken; me-

dial posterior depression present between tubercles; ap-

erture moderately wide, curved strongly posteriorly to-

ward columella: outer lip extending posteriorly beyond

inner lip; outer lip with 17 short simple teeth: columella

with about 13 short weak simple teeth, fading posteriorly;

denticular interstices smooth; terminal ridge not visible;

fossula narrow, mostly covered; siphonal banal long, bor-

dered by spatulate extensions of columella and outer lip;

terminal canals moderately extended, very weakly

rimmed.

Comparison: The new species is most similar to Wood-

ring’s (1959) hypotype of Cypraea (Muracypraea) henek-

eni |= Muracypraea mus isthmica (Schilder, 1927)],

USNM 562603 (pl. 31, figs. 6-7) from the upper Mio-

cene Gatun Formation of Panama. Muracypraea or-

menoi has fewer columellar teeth, a less inflated col-

umella, deeper terminal canals, and is not as highly

arched as M. mus isthmica. Color pattern aside, when

compared to several specimens of the Recent M. mus

bicornis (Sowerby, 1870) [LACM 90-47.1 (LACM Mal-

acology Bie see also Lorenz and Hubert, 2000, pl.

IL, figs. 11, 16, 19], we note a striking similarity in overall

morphology between M. mus bicornis and the new spe-

cies. However, the new species has a more inflated col-

umella, fewer columellar teeth, and deeper terminal ca-

nals than M. mus bicornis. Muracypraea angustirima

(Spieker, 1922), the other Peruvian muracypraeid spe-

cies, from the early to middle Miocene Zorritos Forma-

tion of Tumbés Department, northern Peru, lacks the

dorsal tubercules, has a straighter aperture, has more

columellar teeth, and has a more inflated columella than

the new species.

Discussion: Muracypraea ormenoi is the first cypraeid

reported from the Pisco Basin. Although post-burial pro-

cesses have damaged the mid-dorsal and the posterior

dorsal surfaces and the anterior columellar terminal

ridge of the holotype of the new species, preservation is

adequate for unequivocal generic assignment. The non-

type specimen is a poorly preserved specimen consisting

of three fragments, including labral and columellar lips

and several Teseoal whorls within matrix. Some mollusks

associated with M. ormenoi in Peru range southward to

the Miocene Navidad Basin of central Chile (Table 1;

DeVries and Frassinetti, 2003). Other species associated

with M. ormenoi range northward to the Talara Basin of

northern Peru, situated at the same low latitudes where

warm sea surface temperatures help define the modern

Panamic Faunal Province. Despite the warm-water en-

vironment indicated by the southern Peruvian Miocene

faunas, sedimentological indicators including diatoma-

ceous siltstone, synsedimentary horizons of vemepilized

dolomite, and phosphatic concretions (Dunbar et al.,

1990) point to the existence of high primary productivity

along the southern Peruvian margin during the early and

middle Miocene. Then, as now, the impetus for high

primary production and would have been coastal up-

welling. Upwelled waters were probably cooler than sur-

face waters, but temperatures may still have been more

elevated at present, since the ‘world’s oceans were

warmer during the early Miocene, as well (Savin et al.,

1985; Lourens et al., 2004).

Material: §=Muracypraea ormenoi new species is repre-

sented by the moderately well-preserved holotype from

the upper Oligocene to lower Miocene Chilcatay For-

mation and an incomplete specimen from the overlying

lower middle Miocene to upper Pliocene Pisco Forma-

tion. The non- type specimen (LACMIP no. 13070) was

collected by the senior author in June, 1987, near Cerro

Submarino, east of Yesera de Amara, Ica Department,

Table 1. Mollusks from the Chilcatay and lowermost Pisco formations of southern Peru associated with Muracypraea armenoi new

species and their occurrence in depositional basins to the north and south.

Species

Talara Basin Northern Peru

Navidad Basin Central Chile

Acanthina katzi (Fleming, 1972)

Eucrassatella ponderosa (Philippi, 1987)

Ficus distans (Sowerby, 1846)

Glycymeris colchaguensis (Hupé, 1954)

Glycymeris ibariformis (Frassinetti and Covacevich, 1984)

Miltha vidali (Philippi, 1887)

Olivancellaria claneophila (Duclos, 1835)

Testallium cepa (Sowerby, 1846)

Architectonica karsteni (Rutsch, 1934)

Chionopsis sp.

Conus spp

Dosinia sp

Gonyscyon sp.

Phyllonotus sp.

Terebra sp

Turritella woodsi (Lisson, 1925)

Turritella infracarinata (Gryzbowski, 1899)

hope pe ee be oe be Oe

xs KK KK KK KB

T. DeVries et al., 2006

Page 105

Peru (14°34'26"S, 75°40'30’W) [DV 553-1

loc. 17615]

Type Material: Holotype LACMIP no. 8197, 64.9

mim in length, 48.3 mm in width, 31.9 mm in height.

Collected in October 2004 by Manuel Ormeno, Ocucaje,

Peru.

, = LACMIP

Type Locality: LACMIP loc. 17783, between Yesera

de Amara and the Rio Ica, less than one km north of

Cerros Las Tres Pyramides [herein designated Ullujalla

West| (about 14°34'50" S, 75°38'40" W; Lomitas 1:

LOO,OOO quadrangle), Ica Department, Sothern Peru,

upper Oligocene to lower Miocene Chilcatay Formation

(Figure 2).

Etymology: Named after Manuel Ormefo who col-

lected the holotype and was one of the principal field

workers employed by Mario Urbina.

ACKNOWLEDGMENTS

We wish to thank Rodolfo Salas of the Universidad Na-

cional Mayor de San Marcos, Museo de Historia Natural,

Laboratorio de Vertebrados, Lima, Peru, and Dr. Niels

Valencia, director of the museum, for their support of

invertebrate paleontology in Peru. We also thank James

H. McLean and Angel Valdés (LACM Malacology),

LouElla Saul (LACMIP), and Richard L. Squires (C Cali-

fornia State University, Northridge, Geological Sciences)

for reviewing early drafts of the manuscript and offering

as suggestions. The evaluations of Charles 2 Pow-

ell, (U.S. Geological Survey, Menlo Park, CA) and

ae J. Stanton (LACMIP) ) greatly canes this pa-

per. Donald W. McNamee (LAC M Research Library)

assisted the second author in locating several obscure

references. Additional thanks to Angel Valdés for his

generous assistance with the manipulation of digital im-

ages.

LITERATURE CITED

Adams, G. I. 1909. An outline review of the geology of Peru.

Annual Report of the Board of Regents of the Smithsonian

Institution for 1908: 385-430, figs. 1-12, pls. 1-5.

DeVries, T. J. 1998. Oligocene deposition and Cenozoic se-

quence boundaries in the Pisco Basin (Peru). Journal of

South American Earth Sciences 11: 217-231.

DeVries, T. J. 2001. Molluscan evidence for an Oligocene-

Miocene age of ‘Paracas’ beds in southern Peru. Boletin

de la Sociedad Geoldégica del Perti 92: 57-65.

DeVries, T. J. and D. Frassinetti. 2003. Range extensions and

biogeographic implications of Chilean Neoge ne mollusks

found in Peru. Boletin del Museo Nacional de Historia

Natural, Chile 52: 141-157.

DeVries, T. J. and H. Schrader. 1997. Middle Miocene marine

sediments in the Pisco Basin (Peru). Boletin de la So-

ciedad Geoldgica del Perit 87: 1-13.

Dunbar, R. B., R. C. Marty, and P. A. Baker. 1990. Cenozoic

marine sedimentation in the Sechura and Pisco basins,

Peru. Palaeogeography, Palaeoclimatology, Palaeoecology

77: 235-261.

Groves, L. T. 1997. A review of cypraeiform gastropods from

Neogene strata of northwestern Ecuador, with the de-

scription of two new species. Tulane Studies in Geology

and Paleontology 30; 147-158.

Groves, L. T. 1998. The cypraeid genus Murac ypraca Wood-

ring, 1957 (Mollusca: Gastropoda) in the Caribbean and

adjacent regions: How many pe cies are there and what is

M. henekeni (Sowerby, 1850)? PaleoBios 1S ( (supplement

to 3): 3.

Kay, E. A. 1996. Evolutionary radiations in the Cypraeidae.

Chapter 1. In: Taylor, J. D. (ed.), Origin and evolutionary

radiation of the Mollusca. Oxford University Press, Lon-

don, pp. 211-220.

Linnaeus, C. 1758. Systema naturae per regna tria naturae.

Edito decima, reformata. Regnum animale 1, Holmiae,

824 pp.

Lorenz, F. and A. Hubert. 2000. A guide to worldwide cowries.

2"" edition. ConchBooks, Hackenheim, 584 pp:

Lourens, L., F. Hilgen, N.J. Shackleton, J. Laskar, and D.

Wilson. 2004. The Neogene period. Chapter 21. In: Grad-

stein, F. M., J. G. Ogg, and A. G. Smith (eds.) A geologic

timescale 2004. Cambridge University Press, Cambridge,

pp. 409-440.

Macharé, J. and I. Fourtanier. 1987. Datation des formations

Tertiaires du Bassin de Pisco (Pérou) a partir

dassociations de diatomées. Comptes Rendus de

lAcadémie des Sciences, Paris, Série 2, 305(5); 407-412.

Marks, J. G. 1951. Miocene stratigraphy and paleontology of

southwestern Ecuador, Bulletins of American Paleoniel-

ogy 33(139): 277-433.

Olsson, A. A. 1964. Neogene mollusks from northwestern Ec-

uador. Paleontological Research Institution, Ithaca, 256

pp. :

Pilsbry, H. A. and A. A. Olsson. 1941. A Pliocene fauna from

western Ecuador. Proceedings of the Academy of Natural

Sciences of Philadelphia 93: 1-79.

Rafinesque, C.S. 1815, Analyse de la nature, ou tableau de

univers et des corps organisés. Palermo, 224 pp.

Savin, S. M., L. Abel, E. Barrera, D. Hodell, J. P. Kennett, M.

Murphy, G. keller, J. Killingley, and E. Vincent. 1985.

The evolution of Miocene surface and near-surface marine

temperatures: Oxygen isotopic evidence. In: Kennett, J.P.

(ed.) The Miocene ocean: Paleoceanography and biogeog-

raphy. Geological Society of America Memoir 163: 49-82.

Schilder, F. A. 1927. Revision der Cypraeacea (Moll., Gastr.).

Archiv fiir Naturgeschichte 91A(10): 1-171.

Schilder, M. and F. A. Schilder. 1971. A catalogue of fossil and

living cowries. Institut Royal des Sciences Naturelles de

Be ‘Igique, Mémoire 85: 1-246.

Sowerby, G. B., IL. 1850. Descriptions of new species of fossil

shells found by J. S. Heniker Esq. Proceedings of the

Geological Society of London, Quarterly Journal 6: 44-53,

vs 9-10.

Sowerby, G. B., IL. 1870. Monograph of the genus Cypraea.

Thesaurus Cc onchyliorum 4. London, 58 pp., 37 pls.

Spieker, E. M. 192: 2. The paleontology of the a Forma-

tion of the northern Peruvian oil fields. The Johns Hopkins

University Studies in Geology 3: 1-197, pls. 1-10.

Woodring, W. P. 1957. Muracypraea, new subgenus of

Cypraca. The Nautilus 70: 88-90,

Woodring, W. P. 1959. Geology and paleontology of Canal

Zone and adjoining parts of Panama. De scription of ne r-

tiary mollusks (Gastropods: Vermetidae to Thaididae

U.S. Geological Survey an Paper 306-B:II] + ven

239.

THE NAUTILUS 120(3):106-111, 2006

Page 106

The status of unionid and dreissenid mussels in northwestern

Pennsylvania inland lakes

K. J. Butkas' and M. L. Ostrofsky”

Allegheny College

Biology Dep: iurtment

Meadville, PA 16335 USA

[email protected]

ABSTRACT

A survey of the native unionid fauna of the eight natural lakes

in the glaciated region of northwestemm Pe nnsylvania suggests

that the documented species richness of regional lotic habitats

cannot be extended to lentic habitats. Richness r: anged from 0

to 5 species per lake, and population densities (<0.03 individu-

als/m~) are low. The most frequent and abundant species were

Pyganodon grandis and Utterbackia imbecillis, each occurring

in five lakes, and making up 50% and 15% respectively of all

unionids collected. Two of the lakes have been successfully

colonized by Driessena polymorpha, with evidence of an un-

successful introduction in a third. In Sandy Lake, 75% of the

unionids were fouled by Dreissena, with an average density of

271 Dreissena/unionid. Although none of the seven species of

native unionids found in this survey are regionally endangered,

the prognosis for lake populations is poor.

Additional Keywords: Unionids, zebra mussels,

lakes, Corbicula, Dreissena.

Pennsylvania

INTRODUCTION

With a historical richness of 292 species of Unionidae

and five species of Margaritiferidae, North America’s

freshwater mussel richaees ¢ (superfamily Unionoidea) is

five times greater than that of any of the other continents

(Bogan, 1993: Williams et al., 1993: Lydeard et al., 2004).

However, as part of a worldwide pattern of decline in

many non-marine mollusks, this fauna is declining rap-

idly and much faster than the continent’s mammal or

bird fauna (Williams et al., 1993: Lydeard et al., 2004).

The National Heritage Network ae 202 species that are

presumed extinct, imperiled or vulnerable (Lydeard,

2004), and the Endangered pase: Act recognizes that

13 species in the U.S. have gone extinct and that another

70 are threatened or endangered (U.S. Fish and Wildlife

Service, 2005).

1 1 1 . .

Present address: Center for Limnology, University of Wiscon-

sin, Madison

2 orresponding author mostrofs@allegheny edu

Potential reasons for these declines include siltation;

release of toxic effluents from industrial, municipal, and

nonpoint sources; channel modification; damming; heavy

metal and pesticide bioaccumulation; anoxia due to an-

thropogenic eutrophication; a history of commercial ex-

ploitation; and the introduction of invasive species, par-

ticularly the zebra and quagga mussels (Dreissena poly-

morpha (Pallas, 1771) and Dreissena bugensis

(Andrusov, 1897) (Nalepa et al., 1991; Bogan, 1993; Gou-

dreau et al., 1993; Gillis and Mackie, 1994; Nalepa, 1994;

Schloesser and Nalepa, 1994; Ricciardi et al,. 1996;

Schloesser et al., 1998; Poole and Downing, 2004).

It is unclear how this loss of unionid richness and

biomass will affect ecosystem function; burrowing bi-

valves have the potential to play important roles in many

benthic and pelagic ey but these roles have gone

largely understudied (Vaughn and Hakenkamp, 2001).

However, rapid declines have focused recent attention

on many facets of mussel biology. Researchers are work-

ing to better understand unionid life histories, distribu-

tion patterns, and taxonomy so that conservation efforts

may be more effective. Part of this work involves taking

inventory of current mussel diversity and distribution in

various locales and focusing attention on conservation

“hotspots”—places that are like ‘ly to harbor relatively high

diversity.

The Pennsylvania portion of the Ohio River watershed

is one of these “hotspots.” Before the recent wave of

extinctions there was a relatively rich fauna of 53 species

in this drainage (Bogan, 1992). Twelve species have been

extirpated from the area, but 41 remain, including two

federally endangered species Epioblasma torulosa rangi-

ana (Lea, 1838) and Pleurobema clava (Lamarck, 1819)

(Pennsylvania Biological Survey, 2003). Because of this

relatively high niékne ss, unionid populations in the lotic

waters of Pennsylvania's Ohio River drain: ie have re-

ceived conside rable attention from such groups as the

French Creek Project (Meadville, Pennsylvania), The

Nature Conservancy, and the Western Pe nnsylvania

Conservancy (Pittsburgh, Pennsylvania).

In contrast, the lentic waterbodies within Pe mnsylva-

nia’s Ohio River drainage have received much less atten-

K. J. Butkas and M. L. Ostrotsky, 2006

Page 107

tion. It is possible that these lakes harbor some of the

region’s mussel diversity since several of them flow into

French Creek, a pi articul: uly unionid-rich tributary of the

Allegheny and Ohio Rivers. French Creek he urbors 27

species of unionids, including Epioblasma torulosa ran-

giana and Pleuroblema clava.

These lakes are also at risk for dreissenid colonization

because of active recreational boat traffic in the area, and

close proximity to Lake Erie and to each other. If suc-

cessfully colonized by dreissenids, these lakes could serve

as reservoirs from which dreissenids could then travel to

other waterbodies either by going downstream via pas-

sive means or by going overland via human transport.

Thus, successful dreissenid colonization could negatively

affect regional unionid diversity as it did in many other

drainage »s (Gillis and Mackie, 1994: Schloesser and

Nalepa, 1994; Ricciardi et al., 1996; Schloesser et al.,

1998).

In light of the potential threat of dreissenid coloniza-

tion of these lakes and the current and historical levels of

regional unionid diversity, we sought to accomplish the

following goals: 1) Provide a benchmark assessment of

unionid richness me density in the natural lakes of

northwestern Pennsylvania and 2) Determine the extent

of successful dreissenid and any other invasive mollusk

colonization.

STUDY AREA

Natural lakes in northwestern Pennsylvania are limited to

a relatively small glaciated region (approx. 9200 km?) that

includes all of Pirie. @ canard, and Mercer counties, and

small portions of adjacent counties. Within this region

there exist § kettle Iskes at elevations between 300 and

400 m above sea level (Figure 1). The lakes range in size

from 6 to 378 ha, and are all between 20 and 80 km from

the south shoreline of Lake Erie. However, all discharge

to the south into the Ohio-Mississippi river drainage.

Calcareous glacial deposits dominate the surface geol-

ogy, and dominant land cover is mixed forest and agri-

culture. As a consequence, the lakes have moderately

hard water, are mesotrophic to eutrophic, and undergo

periodic deep-water anoxia during the summer months.

All are headwater lakes, and most are draine d by perma-

nent streams that allow for easy dispersal of aquatic or-

ganisms. The single exception is Edinboro Lake that was

enlarged by the construction of a 3 m hydraulic dam in

the 19" century. Discharges from Canadohta and Con-

neaut lakes are also regulated to prevent ice damage to

docks, but the outlets are unimpeded between October

and May. Physical and chemical characteristics of the

lakes are shown in Tables 1 and 2.

MATERIALS AND METHODS

A census of between 15 and 20 quadrats (7.61.5 m) took

place in each of the eight lakes between August and

October 2002. Three to four locations were chosen in

Lake Erie

~Allegheny River

Figure 1. Study area showing the location of lakes and major

streams. L. Camilones Lake; 2. Conneaut Lake; 3. Cry stal Lake:

4. Edinboro Lake; 5. Lake LeBoeuf; 6. Lake Pleasant; 7. Sandy

Lake; 8. Sugar Lake.

each lake to capture nearshore physical habitat variation.

For example, if a lake had sandy, rocky, and weedy areas,

all three were sampled. At each location, five quadrats

were established—one from each of the following depth

intervals: 0-0.6m, 0.7-1.2m, 1.3-1.S8m, 1.9-2.4m, and

2.5-3.0m. In most cases, the nearshore substrate condi-

tions gave way to finer substrata as depth increased.

Quadrats were delineated with nylon rope strung be-

tween PVC pipe “stakes” that were pushed vertically into

the sediments or placed flat on impenetrable substrata

such as rock. Using the nylon ropes as visual and tactile

guides, a diver manually probed the substrate to a depth

of about 5 cm for bivalves. Each quadrat was searched

twice.

In quadrats with exceptionally high densities of dreis-

senids a 1.5x0.6 m sub-quadrat was delimited: For dre-

issenids, we tallied the number of live individuals. For

unionids, we tallied both live and recently dead individu-

als of all species. Recently dead unionids were those

whose valves could be pz aired and identified to species.

Total unionid and dreissenid densities were recorded as

individuals/m*. Where unionids were fouled with dreis-

senids, the number of dreissenids per unionid was also

recorded.

Bivalves were identified to species level using Bogan

(1992), Strayer and Jirka (1997), Parmalee and Bogan

(1998), and C ummings and Mayer (1992). Nomenclature

follows Turgeon et al. (1998).

In addition to quadrats described above, littoral sub-

Page 108

THE NAUTILUS, Vol. 120, No. 3

Table 1. Physical characteristics of northwestern Pennsylvania lakes. (Current bathymetric maps for Lake LeBoeuf and Sandy Lake

are unavailable.)

Drainage basin

Max depth

Mean depth

Lake Location Area (ha) area (km? ) (m) (m) Outflow

Canadohta 41°48.91'N 68 20.3 13.6 5.5 Oil Creek

79°50.28'W

Conneaut 41°37.5'N 378 72.3 19.8 5.6 Conneaut Outlet to French Creek

$0°18.3'W

Crystal 41°33.21'N 6.1 0.28 7.1 3.3 Crooked Creek to Shenango River

: §0°22.14'W

Edinboro 41°52.78'N 167 65.7 9.1 3.4 Conneauttee Creek to French Creek

80°08 .22'W

LeBoeuf 41°55.61'N 32.2 158.6 9.1 — LeBoeuf Creek to French Creek

79°58.96'W

Pleasant 42°00.22'N 93.7 3.6 3.7 6.8 Lake Pleasant Outlet to French Creek

79°53.85'W

Sandy 41°20.71'N 88.5 7.2 114 — Unnamed stream to Sandy Creek

80°06.43/W

Sugar 41°33.94'N 31 56.4 5.5 3.2 Lake Creek to French Creek

: 79°56.75'W

habitats less than 1.8 m in depth were further sampled

for benthic macroinvertebrates. This was done to im-

prove the likelihood of finding invasive mollusks at low

densities or with small body sizes. We sampled in sandy,

rocky, and weedy littoral sub-habitats, although not all of

these sub-habitats were present in each lake. ‘Depending

on the number of sub-habitats present, the total number

of areas sampled per lake ranged from S-17. We used

steel drop boxes (0.125 m~ > and 0.25 m7, on fine/medium-

grained and cobble substrates, respectiv ely) and a corer

(0. 025 m7, where the roots of a species of Nuphar pre-

cluded sampling with the drop boxes). The substrate iso-

lated by drop boxes was swept 6 times with a 900 jum

mesh D-net. Catches from replicate sweeps in each drop

box sample were washed and pooled. Core samples were

taken to a depth of 10 cm and were similarly washed. The

material was sorted in the laboratory from white tr ays.

Any invasive mollusks were identified using the sources

mentioned above and Jokine n (1992). Densities were ex-

pressed as individuals/m> substrate.

Voucher specimens of the species examined in this

study are deposited in the Section of Mollusks at the

Came »cie Museum of Natural History, Pittsburgh, as fol-

lows:

Actinonaias ligamentina: CMNH 74451; Lampsilis

siliquoidea: CMNH_ 74452; Utterbackia imbecilis:

CMNH 74453; Pyganodon grandis: CMNH 74454; Las-

migona complanata: CMNH 74455; Ligumia recta:

CMNH 74456.

RESULTS

NATIVE UNIONIDS

A survey of the eight natural lakes resulted in a list of

). Pyganodon

seven species of native unionids (Table 3

grandis (Say, 1529) was both the most broadly distrib-

uted, occurring in 5 of the lakes, and the most abundant,

making up half of the individuals collected in all lakes. In

New York State, there has been difficulty surrounding

the taxonomy and identification of the closely related

species Pyganodon cataracta, P. grandis, and P. lacustris

(Strayer and Jirka 1997). However, according to Strayer

and Jirka (1997) and Strayer (Institute of Ecosystem

Studies, personal communication, 2003), P. grandis is

the only one of these three species to occur in the Ohio

basin. Therefore, all Pyganodon collected in this study

were identified as P. grandis even if the beak sculpture

was obliterated or damaged and could not be used as a

definitive identifying character. Utterbackia imbecillis

(Say, 1829) was found in five of the lakes, but it made up

less than 15% of all individuals collected. Lampsilis sili-

quoidea (Barnes, 1823) was found in two lakes, and Ligu-

mia recta (Lamarck, 1819), Lasmigona complanata

(Barnes, 1823), Actinonaias ligamentina (Lamarck,

1819), and Amblema plicata (Say, 1817) were each found

in only one lake. Lake Le Boeuf had the highest species

richness with five species. No mussels were found in

Crystal Lake.

In the 5 lakes where live unionids were present, they

were found in low densities ranging from 0.01 individu-

als/m> in Conneaut Lake to only 0.03 individuals/m2 in

Canadohta Lake. In Pleasant and Sugar Lakes only valves

were found. Unionid distributions and densities are

shown for each lake in Tables 3 and 4.

DREISSENIDS AND OTHER INVASIVE MOLLUSKS

All dreissenids in this study were identified as Dreissena

polymorpha based on morphological characteristics out-

lined by Pathy and Mackie (1993) and confirmed by

Mackie (Univ ersity of Gue Iph, person il communication,

2003).

Living zebra mussels (D. polymorpha) were found

only in Edinboro Lake and in Sandy Lake. In Edinboro

K. J. Butkas and M. L. Ostrotsky, 2006

Page 109

Table 2. Chemical characteristics of northwestern Pennsylva-

nia lakes. P = phosphorus.

Spring Carlson's Total Total

total P_ trophic state alkalinity dissolved

Lake (pg/L) index (mg/L) solids (ppm)

Canadohta —_26.9 52 39.0 SO

Conneaut 18.6 46 73.0 120

Crystal 22.8 49 61.5 130

Edinboro 30.1 53 67.0 150

LeBoeuf 47.5 60 13D 110

Pleasant 35.8 56 95.5 140

Sandy 13.6 42 66.5 140

Sugar 41.2 58 37.0 70

Lake, the mean density of D. polymorpha was 52 indi-

viduals/m? (s.e.=48) and the mean density in Sandy Lake

was 442 individuals/m? (s.e.=137). Dreissena polymor-

pha was densely aggregated on scattered submersed

wood, rocks, and on dha shells of native unionids. In

Edinboro Lake, approximately 21% of the live unionids

and paired valves were fouled with D. polymorpha and

an additional 42% had been recently fouled as evidenced

by attached byssal threads. The average density of foul-

ing D. polymorpha in Edinboro Lake was 1.4/unionid

(s.e.=1.1). In Sandy Lake, 78% of the live unionids and

paired valves were fouled with D. polymorpha, and an

additional 22% had byssal threads only. The average den-

sity of oan D. polymorpha here was 271/unionid

=87

The 2 boxes yielded D. polymorpha valves in two

other lakes. In Lake LeBoeuf shells were found in an

artificially sandy area along the northwestern shore and

probably were transported to the lake in beach sand

dredged and imported from Lake Erie. In Canadohta

Lake, a few valves were found near the Pennsylvania

Fish and Boat Commission access point. Prior to this

study, several live D. polymorpha were found at this

same location in 2001. These individuals probably rep-

resent a failed introduction inadvertently brought in by

recreational boaters.

The drop box samples also revealed the invasive Asi-

atic clam, Corbicula fluminea (Miiller, 1774), in Con-

neaut and Sandy Lakes. Like Dreissena polymorpha, the

distributions were highly aggregated with mean densities

in the two lakes of 454 (s.e.=251) and 216 (s.e=87) indi-

. 5

viduals/m”, respectively. In addition, the invasive snail

Cipangopaludina chinensis (Reeve, 1863) was found in

drop box samples from Canadohta Lake and Lake Pleas-

ant.

DISCUSSION

The unionid assemblages in northwestern Pennsylvania's

natural lakes are species-poor and low in density when

compared to the unionid assemblages in the region’s riv-

ers and streams. This stream fauna is also unique in that

it contains endangered species such as Pleuroblema clava

and Epioblasma torulosa rangiana. In contrast, all of the

taxa in our study lakes are ranked globally as “secure,

common” by the Heritage 1 ranking system and none have

been granted special status unde federal law or Penn-

sylvania code (NatureServe, 2005). In terms of within-

state rankings by non-legislative groups, P. grandis, U.

imbecillis, L. siliquoidea, A. ligamentina and L. recta all

have Heritage state ranks ‘of “secure” or “secure/

vulnerable” (NatureServe, 2005) and have no status ac-

cording to the Pennsylvania Biological Survey (2003).

However, L. oe has a Herite we state rank of

“critically imperiled” ( NatureServe, 2005) and is consid-

ered endangered in the state by the Pennsylvania Bio-

logical Survey (2003). Similar ly, A. plicata has a Heritage

state rank * imperiled” (NatureServe, 2005) and is con-

sidered threatened in PA by the Pennsylvania Biological

Survey (2003).

It is difficult to determine if these two taxa of local

concern (L. complanata and A. plicata) or any of the

other taxa we found represent self-sustaining populations

within our study lakes. Downing et al. (1993) found that

in Lac de L’Achigan, Québec, complete fertilization fail-

ure occurs when Elliptio complanata ( (Lightfoot, 1786) is

found at densities less than 10/m?. In our study area, the

density of all living unionids is less than 0.03/m> and

much lower for a given species, thus, successful sexual

reproduction is iinlileshe On the other hand, Strayer et al.

(1981) found similar low densities of Elliptio complanata

(0.032 individuals/m7) in oligotrophic Mirror Lake, and

the size distribution of individuals there suggest regular

recruitment. Frequent hermaphrodism may freltate

species survival in such low-density populations.

Past and present human impacts on unionid fauna of

these lakes are possible since these waterbodies have

Table 3. Native Unionid species recorded from northwest Pennsylvania lakes.

Canadohta Conneaut Crystal Edinboro LeBoeuf Pleasant Sandy Sugar

Pyganodon grandis + + - + + = + =

Utterbackia imbecillis + = 7 7 + + 4 +

Lampsilis siliquoidea = = - = + a £ =

Lasmigona complanata = = = = 4 = _ _

Actinonaias ligamentina ~ - = + = = = =

Ligumia recta = 5 = mn _ _ = =

Amblema plicata 2 = = _ + = = =

Total number of species 2 1 0 3 5 1 3

Page 110

THE NAUTILUS, Vol. 120, No. 3

Table 4. Densities of live unionids and paired valves. All densities are individual per m>. Values in parentheses are standard errors.

Canadohta Conneaut Crystal Edinboro LeBoeuf Pleasant Sandy Sugar

Live unionids 0.030 0.011 0) 0.029 0.023 0) 0.023 0

(O.O11) (0.008) (0.014) (0.018) (0.010)

Paired empty valves 0.017 0) 0 0.080 0.052 0.011 0.020 0.011

(0.012) (0.039) (0.030) (0.008) (0.009) (0.011)

Combined 0.047 0.011 0 0.109 0.075 0.011 0.040 0.011

(0.019) (0.008) (0.039) (0,028) (0.008) (0.014) (0.011)

been subject to a variety of alterations such as cultural

eutrophication, lakeshore development, fish stocking,

water level modification, etc. However, it is difficult to

ascertain whether the current density, richness, and taxo-

nomic composition are typical for small, nutrient-rich

lakes or if they represent the types of declines in rich-

ness/abundance and the alterations in taxonomic compo-

sition that have occurred in many unionid assemblages

across the continent (e.g., } Nalepa et al., 1991 on pre-

Dreissena unionid declines in Lake Erie). Unionid den-

sities in other small lakes (e.g., small Adirondack lakes,

personal observation) can be much higher than found in

our study area, but richness may naturally be low. For

example, Strayer and Jirka (1997) found that small lakes

in New York State usually contain five species or fewer.

While we have some anecdotal basis for comparison, we

lack models that predict unionid species’ richness or

abundance for lentic systems. Predictors of lentic

unionid richness that are potentially worth exploring in-

clude fish species’ richness and the area of oxygenated

lake bottom (see Watters (1992) for an exploration of

similar predictors in lotic systems).

Another way to assess the current status of unionidae

in PA’s lakes would be to draw comparisons between past

and present. We searched for ea records of the

freshwater mussel fauna in these lakes from the Cleve-

land Museum of Natural History (Cleveland, Ohio), the

Carnegie Museum of Natural History (Pittsburgh, PA),

and Foti documents by key mussel collectors, primarily

A.E. Ortmann, who surveyed f for the Camegie Museum

in the early 20° " century. Specific documents searched

included Ortmann ( (unpublishe ad, 1909, 1911, 1919).

These efforts yielded only species lists in Ortmann (un-

published), a few specimens in the physical collections at

the Carnegie Museum, and a few records in the Car-

negie’s electronic database. These bits of information

summed to an early Q¢th century, anecdotal account for

Conneaut Lake and Edinboro Lake (then called “Con-

neauttee” Lake in Ortmann, unpublished). Thus, overall,

the dearth of quantitative and taxonomically clear infor-

mation on the past mussel fauna of our study area makes

inferences regarding human impacts spe ‘culative at best.

In any case, whether due to natural factors or human

impacts, unionid assemblages in the clacial lakes of

northwestern, PA are currently low in density and low in

species richness and the establishment of zebra mussels

is likely to cause density and species richness to become

even lower. The average number of fouling zebra mus-

sels/unionid specimen in Sandy Lake is well over 100 and

models by Ricciardi et al. (1995) predict that this degree

of fouling will result in greater than 90% unionid mor-

tality. W fietens the average number of fouling Dreissena

polymorpha/unionid specimen was only 1.4 in Edinboro

Lake, this number is Dera not an accurate indicator

of the overall degree of fouling. Winter drawdown has

been used at Edinboro Lake to manage dreissenid popu-

lations (Grazio and Montz, 2002) and the lake was drawn

down approximately 1.5 m during the winter prior to this

study. Thus many of the zebra ‘aqussells that were fouling

unionids in depths shallower than 1.5 m were killed due

to exposure, leaving only byssal threads on 42% of our

specimens (22% of the specimens were fouled by byssal

threads only in Sandy Lake). However, below the draw-

down level, zebra mussels persist in field densities up to

727 individuals/m> and thus are likely to keep fouling

unionids even if periodic drawdown is successful in re-

ducing D. polymorpha in shallow depths.

Because the unionids in these lakes possibly do not

represent self-sustaining populations, it is unlikely

that D. polymorpha- ‘nduced losses within these water-

bodies will nee to any further decrease in richness

or abundance on a regional ‘scale. However, the potential

spread of D. polymorpha from the lentic waterbodies of

northwestern PA to nearby unionid-rich lotic systems is a

potential threat to regional diversity. In addition, while

not necessarily a diet to unionid populations, the pres-

ence of Corbicula fluminea in Conneaut Lake and Sandy

Lake (Strayer, 1999) and the invasive snail C ipangopalu-

dina chinensis in Lake Pleasant and Canadohta Lake

highlights the need for continued public education as to

the causes and potential impacts of the spread of invasive

species.

ACKNOWLEDGMENTS

We would like to thank T. Pearce for generous access to

the unionid collections of the Carnegie Museum, D.

Strayer and G. Mackie for confirming our identifications,

A. McMillen for her capable field assistance and Penn-

sylvania Sea Grant for financial support.

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THE NAUTILUS 120(3):112-115, 2006

Page 112

Replacement names and type material: examples from Hertlein

and Strong (1940-1951) and Keen (1958)

Eugene V. Coan! Richard E. Petit

Santa Barbara Museum of Natural History P.O. Box 30,

2559 Puesta del Sol Road

Santa Barbara, CA 94105-2936 USA

[email protected]

North Myrtle Beach, S¢

[email protected]

) 29597-0030 USA

ABSTRACT

There is a sharp distinction between replacing a preoccupied

name and describing a new species that is a synonym of such a

name. We cite some examples of where this distinction has

been misinterpreted with regard to type material of Hertlein

and Strong and Keen taxa.

INTRODUCTION

The purpose of this paper is to establish the status of type

material for some eastern Pacific Mollusca. The taxa dis-

cussed were re-catalogued, with presumed loss of their

status as primary types, occasioned by what we consider

to be a misunderstanding of the provisions of the Inter-

national Code of Zoolovical Nomenclature (International

Comninission on Zoological Nomenclature, herein ICZN)

Article 72.7 (ICZN, 1999) and its predecessors in prior

editions of the Code (ICZN, 1961; 1964: Article 72d;

ICZN 1985: Article 72e). These article »s treat the * ‘types

of replacement nominal species” (1961, 1964), “name-

bearing types of nominal species-group taxa bearing re-

placement names” (1985), and “name bearing types of

nominal spe cies- orp taxa denoted by re pl icement

names an nova)” (1999). The four re of the

Code have minor mente in wording, but their intent

and effect are identical. The version now in effect is:

72.7. Name-bearing types of nominal species-

group taxa denoted by replacement names

(nomina nova). [f an author proposes a new spe-

cies-group name expressly as a replacement (a

nomen novum) for an earlier available one, then

the two names are objective synonyms; both of the

nominal taxa have the same name-bearing type de-

spite any simultaneous restriction or application of

the new replacement name (nomen novum) to

particular specimens or any contrary designation

i .

Research Associate

of type, or any different taxonomic usage of the

new replacement name. [emphasis supplied]

The key provision of this Article in all three editions of

the Code, with only minor differences in wording, applies

only to names proposed expressly as replacement

names. There is an important distinction between pro-

posing a replacement name and re-describing a species

for which there is no available name that can be used,

and it is the failure to appreciate that distinction that

necessitates this paper.

Ifa species name is found to be preoccupied and there

is no available synonym, a worker discovering that fact

has two options:

(1) The taxon can simply be renamed (nomen novum)

with a simple statement such as “this name is proposed to

replace the preoccupied ___.” Under this procedure

nothing changes except that the replaced name becomes

an objectiv e synonym of the new name. The type mate-

rial is identical for both the old and the replacement

taxon, regardless of any contradictory statements.

(2) The taxon can be described as if it were a new spe-

cies, with types and type locality designated, with the

former name that is a junior homonym tie coming a syn-

onym. In this case, the new taxon has the status of anew

species with its own type material and locality, and the

older name exists as a subjective synonym with its own

type material, if any, and type locality. It is relevant only

if at some future date it is discovered to not be conspe-

cific with the newly proposed taxon.

There can be complications inherent in the first op-

tion, especially if the name to be replaced is from the old

literature, and the location or even existence of type ma-

terial is uncertain. Proposing replacement names for

nomina for which there is incomplete information and/or

no known type material was recently addressed by

Rosenberg and Petit (2001). Description of a species as

new provides accessible type material and other data.

This procedure is often used when monographers wish to

establish a firmer basis for the species involved than it

would be possible by simple substitution for a new name.

E. V. Coan and R. E. Petit, 2006

The distinction between these two procedures is clear

but, as with much systematic nomenclature, it can be-

come clouded. Indeed, none of the four versions of the

Code are completely clear for cases in which both the

terminology (a) “new species” and (b) “replacement

hame” or “new name” are simultane ously present. How-

ever, the phrase “expressly as a replace ment name” is a

c leap demarcation, especially the word “expressly” that

appears in all versions of the applicable Article.

In each of the examples cited below, type material was

set aside by the authors involved and catalogued as such,

at the California Academy of Sciences for the Hertlein

and Strong taxa and at Sti anford Unive rsity for the Keen

taxa, with the latter subsequently transferred to the Cali-

fornia Academy of Sciences [as CAS below] with the rest

of the Stanford collection. Then, later, probably in the

1980s, all of the supposed type material was re-

catalogued as non-type specimens.

Hertlein and Strong (1940-1951) and Keen (1958), i

part motivated by the lack of type material on the W, est

Coast and the difficulty either of visiting European or

East Coast institutions or obtaining loans or photographs

of type material, utilized a modification of homonym re-

naming. While stating that the motivation for their taxo-

nomic actions was the presence of a situation of hom-

onymy, they clearly stated their desire to propose new

species with new type material. In other words, in the

terminology of the Code published a few years later, they

were trying to avoid having new taxa being interpreted

merely as Sato nt names. Although iee actions

took plac e before the publication of the Feet of the “mod-

ern” Codes, the current Code makes clear that it and only

it is the governing authority (ICZN 1999: Article 86.3).

The nomina of concern in this paper, and our judgment

as to their type material, are:

A. HERTLEIN AND STRONG PAPERS

In their research on Panamic mollusks, Hertlein and

Strong (1940-1951) found names they believed to be

homonyms. In some cases discussed below, the names

involved were mistakenly thought to be primary or sec-

ondary homonyms, raising dierent issues that are not

discussed in detail here. The { irst two names below were

proposed in a short note in The Nautilus written for the

sole purpose of replacing the preoccupied names.

(1) Lucina undatoides Hertlein and Strong, 1945: 105.

“The species named Lucina undata by Carpenter

requires a new name because of the prior use of that

combination of names by Lamarck, 1819. The name Lu-

cina undatoides is here proposed for the species de-

scribed by Carpenter. Type in the California Academy of

Sciences paleontology type collection, from Isthmus

Cove, Espiritu Santo Island, Gulf of California.”

The simultaneous designation of new type material

and locality does not saad. and their type material does

not have type status. It will elsewhere be demonstrated

that this was not really a homonym in the first place and

return to use of Carpenter’s name for the eastern Pacific

taxon will be recommended (Coan and Valentich-Scott,

2006, in preparation).

Tellina liana Hertlein and Strong, 1945: 105,

“The combination of names, Tellina panamensis used

by Li... had already been used by Philippi in 1848. The

name Tellina liana is here proposed for the species de-

scribed by Li. Type in the California Academy of Sci-

ences pale ontology type collection, dre dge doff

Meanguera Island, E ‘| Salvador, in the Gulf of Fonseca,

in 16 thoms?

As in the previous case, their new type material des-

ignation has no status as such. Moreover, the replace-

ment name was unnecessary, because Li’s species was

originally spelled T. panamanensis. (Li's species and

Hertlein and Strong’s replacement are both synonyms of

Tellina eburnea Hanley, 1844 [Coan and Valentich-Scott,

in prep.]).

In a series of papers on west Mexican mollusks pub-

lished in Zoologica, Hertlein and Strong treated preoc-

cupied taxa differently. The following seven examples

(3-9) are different and in some cases more complex.

Headings are as published.

(3) Arca (Arca) fernandezensis Hertein and Strong,

nom. nov.—(1943: 154)

This name was proposed as a replacement name for

Arca angulata King and Broderip, 1532, non “Meuschen,

1787” [an unavailable, non-binominal work]. We have

not located an available use of A. angulata prior to 1832.

concluding that this replacement was unnecessary, and

that workers should return to the use of the King and

Broderip name. It must be noted here that thie new

name is followed by the abbreviation “nom. nov.”

(nomen novum), it is clearly a replacement name, and

the authors did not attempt to designate new type ma-

terial.

In six other cases, they referred to the new nomina as

new species in addition to designating new type material.

) Cardium (Americardia) guanacastense Hertlein and

Strong, sp. nov.—(1947: 140)

Thei ir entry is headed “sp. nov.” as shown. The shell is

formally de scribed in detail, with comparison and distri-

bution, with new ty spe material and a new type locality. In

the discussion the authors stated “Sowe rby’s combination

of names, Cardium planicostatum, has been used earlier

by Sedgwick and Murchison; therefore a new name is

re quired for Cardium planicostatum of Sowerby and the

name Cardium ee astense, based upon a ty pe from

Culebra Bay, Costa Rica, is here proposed.”

The curatorial staff at the California Acade my of Sci-

ences decided that the type material had no type status,

a determination here considered to be incorrect. How-

ever, as shown elsewhere (Coan, 2002), the homonymy

never existed and the correct name to use is Americardia

planicostatum (G. B. Sowerby I, 1833)

Page 114

THE NAUTILUS, Vol. 120, No. 3

(5) Pitar (Pitarella) mexicanus Hertlein and Strong, sp.

nov.—(1948: 171)

This was also shown as a new species, with a complete

description, comparisons, distribution and new type ma-

terial and locality. In their discussion, the authors stated:

“The species described as Cyther rea lenis by Conrad

[1848] was referred to the genus “Pitaria” [= Pitar] by

K. V. W. Palmer [1927]. This: necessitates a new name for

the west American species described under the name of

Pitar lenis by Pilsbry and Lowe [1932] and the name

Pitar mexicanus, based on a specimen off Maldanado

Point, Mexico, is proposed.” The CAS staff determined

that their type material had no type status.

(6) Tellina (Moerella) recurvata Hertlein and Strong, sp.

nov.—(1949: 71)

Listed as a new species as shown, with a complete

description, comparisons, distribution and with new type

material and locality. In this case, they only cited ge

(Angulus) recurva ‘Dall. 1900, non Deshayes, 1855,

their synonymy. They did not use the words “new name”,

“replacement name”, or “nom. nov.” in their treatment.

The CAS staff later determined that their type material

has no type status.

(7) Tellina (Merisca) proclivis Hertlein and Strong, sp.

nov.—(1949: 83)

Treated as a new species, with a complete description,

comparisons, distribution, and with new type material

and locality. They said, “The use of the combination of

names, Tellina declivis by Conrad, 1534, makes it nec-

essary to propose a new name which is based on a new

type specimen for the west American shell described

under that name by Sowerby in 1868 which is here

named Tellina proclivis.” (Tellina declivis G. B. Sowerby

I, 1868, proved to be a Macoma from another province;

its actual provenance and identity with other taxa is as yet

uncertain, and it is for the time being considered a

nomen dubium.)

Hertlein (1968) later decided that this was the case of

a replacement name, and he then named the eastern

Pacific taxon Tellina ulloana based on the above type

material. Depending on one’s interpretation of this situ-

ation, the eastern Pacific taxon may now have two names,

with Sowerby’s unknown T. declivis remaining un-

renamed, and it is that position that we endorse. This is

an example of the dangers in simply renaming a junior

homonym unless one can be positive about what exactly

is being renamed! The CAS collection at present indi-

cates chat the type material of T. proclivis has no type

status under that name.

8) Latirus mediamericanus Hertlein and Strong,

i L951: SO)

Listed as a new species with a complete description,

comparisons, distribution, and with new type material

ancl type locality. They said, “The name Turbinella cas-

fanea was first proposed by Gray in 1839 for a shell

which Melvill later referred to the synonymy of Leuco-

sp.

zonia cingulifera Lamarck, . . . It therefore becomes nec-

essary to propose a new name for Reeve’s Turbinella

castanea and the name Latirus mediamericana is here

proposed.” The staff at the CAS determined that their

type material has no type status.

(9) Muricopsis zeteki Hertlein and Strong,

(1951: 85)

Listed as a new species, with a complete description,

comparisons, distribution, and with new type material

and locality. The chresonymy [a synonymy that includes

misuses of names as well as true synonyms] begins with

Murex aculeatus Wood, 1828, which is preoccupied by

M. aculeatus Lamarck, 1822. Next listed is M. dubius

Sowerby, 1841, which is a replacement name for M. ac-

uleatus Lamarck. Hertlein and Strong stated that “The

name Murex dubius which was applied to this species by

Sowerby is not valid because that combination of names

had already been applied to a different species by

Dillwyn in 1817. We base the new name, Muricopsis

zeteki, upon a Soa collected by Dr. James Zetek at

Panama City, Panama. Nowhere is it stated that Muri-

copsis zeteki is a replacement name. However, the staff

at the CAS determined that their type material has no

type status.

sp. nov.—

B. KEEN (1958)

Keen (1958) published a paper in which she named sev-

eral new species because of underlying homonymy. In

the abstract to that paper, it is stated thats ‘Six homony-

mous names are cited, of which four are replaced.” In the

Introduction to the paper, she said: “During the prepa-

ration of a handbook on tropical West American marine

mollusks, it was perhaps inevitable that I should find a

few homonyms, but I did not expect so many as six.

Therefore, the opportunity has been taken here to select

new type specimens by treating the replacements not as

mere new names but as new species. The new type speci-

mens are deposited in museums that may be visited with-

out the necessity of a trip abroad.”

Each of her individual descriptions is clearly indicated

as a new species description with the standar d abbrevia-

tion n. sp. following each of the four new names. There

were available synonyms for the other two of the hom-

onyms she unearthed. The taxa are as follows:

(1) Nuculana (Saccella) fastigata Keen, 1. sp.—(1958:

240)

Nucula gibbosa G. B. Sowerby I, 1833, non Fleming,

1828.

2) Adrana cultrata Keen, n. sp.—(1958: 240)

Nucula elongata G. B. Sowerby I, 1833, non Bose,

1801, nec Detrance, 1825.

(3) Macoma (Psanunacoma) elytrum Keen,

244)

Tellina elongata Hanley,

sp. (1958:

1844, non Dillwyn, 1823.

E. V. Coan and R. E. Petit, 2006

Page 115

(4) Crucibulum personatum Keen. n. sp.—(1958: 247)

Calyptraea radiata Broderip, 1834, non Deshayes,

1S30.

For all four of the taxa listed above Keen gave a full

description, type locality, discussion, and designated ho-

lotypes with repository named. However, the curatorial

staff at the CAS interpreted her new type material as

having no type status.

CONCLUSIONS

Our conclusion with regard to the last six examples from

papers by Hertlein and Strong, and the four examples

from Keen (1958) is that fhe clear intention of these

authors was to propose new species. While they did so

because they considered the only names av ailable for

these taxa to be preoccupied homonyms, they attempted

to avoid their new taxa being interpreted as only being

replacement names. The wordings of their attempts to

do so, in the absence of guiding Code language, may have

been awkward in Hertlein and Strong papers, but the

motivation is clear, and Keen’s intent is even more

clearly expressed. Therefore, these taxa are, and should

be interpreted as, new species and not replacement

names, and the type material set aside by the authors

should have type status. The types designated by the

authors and currently in the collections of the CAS are

here regarded as primary types.

As far as known now, this makes a difference on which

name should be used only in the case of Tellina proclivis,

with that becoming the oldest name from the eastern

Pacific taxon, rather than being a replacement name for

a taxon from an unknown province.

We appreciate the advice of Gary Rosenberg and Rii-

diger Bieler on this nomenclatural question.

LITERATURE CITED

Coan, E. V. 2002. Americardia planicostata (G. B. Sowerby I,

1833), an older name returns (Bivalvia: Cardiidae). The

Festivus 34(10): 123-125.

Coan, E. V. and P. H. Valentich-Scott. 2006. Three nomencla-

tural notes on Panamic bivalves. The Festivus {in prepa-

ration}.

Hertlein, L. G. 1968. Tellina ulloana. A new species from

Magdalena Bay, Baja California, Mexico. The Veliger 11:

90.

Hertlein, L. G. and A. McC, Strong. 1943. Eastern Pacific

expeditions of the New York Zoological Society. XXXII.

Mollusks from the west coast of Mexico and Central

America. Part II. New York Zoological Society, Zoologica

28(3): 149-168, pl. 1

Hertlein, L. G. and A. McC. Strong. 1945. Changes in the

nomenclature of two west American marine bivalve mol-

lusks. The Nautilus 58: 105

Hertlein, L. G. and A. McC. Strong. 1947. Eastern Pacific

expeditions of the New York Zoological Society. XXXVI.

Mollusks from the west coast of Mexico and Central

America. Part V. New York Zoological Society, Zoologica

31(4): 129-150, pl. 1

Hertlein, L. G. and A. McC. Strong. 1948. Eastem Pacific

expeditions of the New York Zoological Society. XXXIX.

Mollusks from the west coast of Mexico and Central

America. Part VI. New York Zoological Society, Zoologica

33(4): 163-198, 2 pls.

Hertlein, L. G. and A. M. Strong. 1949. Eastern Pacific expe-

ditions of the New York Zoological Society. XL. Mollusks

from the west coast of Mexico and Central America. Part

VII. New York Zoological Society, Zoologica 34(2): 63-97,

1 pl. -

Hertlein, L. G. and A. M. Strong. 1951. Eastern Pacific expe-

ditions of the New York Zoological Society, XLUI. Mol-

lusks from the west coast of Mexico and Central America.

Part X. New York Zoological Society, Zoologica 36(2): 66—

120, 11 pls.

International Commission on Zoological Nomenclature. 1961.

International Code of Zoological Nomenclature. Interna-

tional Trust for Zoological Nomenclature, London, xvii +

176 pp. :

International Commission on Zoological Nomenclature. 1964,

International Code of Zoological Nomenclature. Second

edition. Intemational Trust for Zoological Nomenclature,

London, xix + 176 pp.

International Commission on Zoological Nomenclature. 1955.

International Code of Zoological Nomenclature. Third

Edition. International Trust for Zoological Nomenclature,

London, xx + 338 pp.

International Commission on Zoological Nomenclature. 1999.

International Code of Zoological Nomenclature. Fourth

Edition. International Trust for Zoological Nomenclature,

London, xxix + 306 pp.

Keen, A. M. 1958. New mollusks from tropical west America.

Bulletins of American Paleontology 38(172): 235-255, pls.

30-31.

Rosenberg, G. and R. E. Petit. 2001. On the unnecessary re-

naming of homonyms in Pyramidellidae. Journal of

Conchology 3 7(3): 245-251.

THE NAUTILUS 120(3):116-117, 2006

Page 116

Book Review

The Mollusks: A Guide to Their Study,

Collection, and Preservation

Sturm, Charles F., Timothy A. Pearce, and Angel Valdés.

2006. The Mollusks: A Guide to Their Study, Collection,

and Preservation. American Malacological Society, Pitts-

burgh, and Universal Publishers, Boca Raton, xii + 445

pp. ISBN 1-58112-930-0 (paperback), 1-58112-931-9 (e-

book); retail price (paperback): $35.95, from http://

universal-publishers.com.

Organizers and editors of books with many contributors

will attest that it is not always easy to bring such projects

to fruition. This may happen for distinct reasons. Differ-

ent contributors will be inclined to allocate different lev-

els of effort into the preparation of their chapters. Such

unevenness may be enhanced when the book in question

does not deal with cutting-edge research or alluring sci-

entific breakthroughs. As a result of dissimilar levels of

commitment from contributors, completion time and the

depth at which subjects are treated may end up varying

between the different chapters of the book.

Successful editors will be patient but persuasive, and

must have a knack for inviting consistency and adding

balance as one chapter flows into the next. In addition, as

I know to be the case with the volume at hand, the

budget for production and distribution may be very lim-

ited from the onset. If all these desiderati and parameters

are brought into the equation, it is clear that Sturm,

Pearce, and Valdés did a remarkable job in pulling to-

gether “The Mollusks”, as Iwill try to show in this review.

The Mollusks aims to replace “How to Study & Collect

Shells,” a guide published in four editions between 1942

and 1974 by the American Malacological Union (AMU;

now American Malacological Society, AMS). The con-

cept for this new version of the AMS cuide was to as-

semble a strong and diverse group of museum profes-

sionals, research malacologists, and serious collectors to

contribute in different areas. As stated in the preface, the

target audience will consist of “amateur and professional

malacologists,” and the book shall “promote the educi a-

tional mission of the American Malacological Society.

Editor Sturm received the green light from the AMS

council to proceed with the organization of the book

during the 2000 annual meeting in San Francisco. In the

few years that passe d, some of the origins il contributors

have left, new techniques in molluscan systematics were

adopted, in particular those for sequencing and analyzing

molecular data; the web, database te schnology, ancl digital

photography grew by orders of magnitude and saw thei

. itus solidified as bonafide researc q tools. The resulting

AMU guide. It con-

volume more than replaces the L974

sists of 445 pages (in contrast to 1O7 pages in 1974) of

A GUIDE TO THEIR

STUDY, COLLECTION,

AND PRESERVATION

THE MOLLUSKS °

® } Edited by ;

C. F. STURM, T. A. PEARCE, AND A. VALDES

\ A PUBLICATION OF THE AMERICAN MALACOLOGICAL SOCIETY

malacological information that may not be found in any

other single publication. Twenty-nine contributors pre-

pared 31 hi ypters covering topics from “Snorkeling and

SCUBA Diving” to * ‘Rearing Terrestrial Gastropoda,”

passing through “Donating Amateur Collections to Mu-

seums” and “Organizations, Meetings, and Malacology”.

In addition to chapters on methods and techniques,

the reader will find specific treatments of the seven main

molluscan clades. There are chapters on “Aplacophora”

(Amélie H. Scheltema), “Monoplacophora” (Clement L.

Counts, IIL), “Polyplacophora” (Enrico Schwabe and An-

reas Wanninger), “Scaphopoda: The Tusk Shells” (Pat-

rick D. Rey olds) and “Cephalopoda” (Frank E. Ander-

son). Gastrop¢ ids are treated in four chapters, “Freshwa-

ter Gastropoda” ( (Robert T. Dillon, Jr.), “Terrestrial

Gastropoda’ (Timothy A. Pearce and Aydin Orstan),

“Rearing Terrestrial oe (Aydin Orstan), and

“Marine Gastropoda” (Daniel L. Geiger), and bivalves in

three (“Unionoida: Freshwater Mussels” (Kevin S$. Cum-

mings and Arthur E. Bogan), “Non-Unionoid Bivalvia”

(Alexei V. Korniushin), and “Marine Bivalvia” (Eugene

V. Coan and Paul Valentich-Scott).

Some contributions may be particularly valuable to

collectors, collection managers, and poloe in gener ral.

“Archival and Curatorial Methods” (Charlie F. Sturm)

discusses a plethora of data on problems ee may arise

when inadequate materials and preparations are involved

in the curation of dry and preserved specimens. Sturm

Book Review, 2006

Page 117

provided a list of materials and suppliers that will prove

handy both to amateur collectors and institutional col-

lection managers. Two chapters on photography, “Digital

Imaging: Flatbed Scanners and Digital Cameras” (F Abio

Moretzsohn) and “ Applied Film Ealeiaohy in System-

atic Malacology” (Daniel L. Geiger), better than ad-

equately cover the topics with examples drawn from the

authors’ obvious experiences. “The Molluscan Litera-

ture: Geographic and Taxonomical Works” (Sturm and

collaborators) offers a brief overview of the malacological

literature in which the main types of taxonomic works are

discussed and a valuable list of regional publications pro-

vided. Other well-researched articles dealing with recent

conceptual and methodological advances are “Comput-

erizing Shell Collections” (Gary Rosenberg), “Cladistics

and Molecular Techniques: A Primer” (David Camp-

bell), and “Issues in Marine Conservation” (Patrick

Baker).

Are there any gaffes or omissions? Certainly, as one

could expect from such a bold proje ct. The resolution of

halftone illustrations (known as “ruling” in typographical

lingo) very coarse, with evident joes of detail. In my

copy of the book, a typo lingers in the title of Daniel

Geiger’s article on “Applie ld [sic] photography ...°; I

read “Monoplacpophora” on page 11S, and “Xenopo-

ridae” on page 120. I miss having access to a subject

index at the end of the volume. Although I realize that

the book is aimed at a very broad audience, | would have

loved to see chapters on applications of electronic mi-

croscopy and georeferencing. At least some of these de-

ficiencies and absences result from the need to keep

production costs within budget, and I can appreciate

that.

Tam not proposing that this is the definitive tome on

malacological techniques and methods because, now

more than ever, techniques and methods change fast,

rendering this type of guide obsolete at the blink of an

eye (think digital imagery and techniques in molecular

systematics, to cite only two ex xamples), However, in

preparation for inevitable obsolence, the editors have

assembled the book in a format that will easily lend itself

to updates. If the editors, contributors, AMS, and the

audience are willing to keep the project alive as de-

manded in fast- evolving times, then this will be the dy-

namic, definitive guide. Until then, log onto the publish-

ers site, get your copy, and encourage your colleagues,

collecting buddies, librarians, and ispaletore acquaintan-

ces to acquire and promote this great-value guide.

José H. Leal

The Bailey-Matthews Shell Museum

P.O. Box Dabs

Sanibel, FL 33957 USA

‘led @dhallcausmuniccve

Sponsored in part by the State of Florida, Department

of State, Division of Cultural Affairs, the Florida Arts

Council and the National Endowment for the Arts

NATIONAL

ENDOWMENT

FOR THE ARTS

5 WHSE

INSTRUCTIONS TO AUTHORS

wii

05822

- S$

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Volume 120. Number 4

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CONTENTS ee aie

John Slapcinsky Paryphantopsis (Gastropoda: Pulmonata: Charopidae) from the Louisiade

Archipelago of New Guinea... 0... eee 119

Gerald E. Walsh Daedalochila lithica and Daedalochila dorfeuilliana (Gastropoda:

Brian F. Coles Polygyridae) in Arkansas, USA: morphology, distribution, and habitat .... 131

Thomas J. DeVries The Neogene history of Prisogaster Mérch, 1550 (Gastropoda:

Turbinidae) in South America 0. ee 139

Marta Dominguez A new species of Hoplodoris Bergh, 1880 (Gastropoda: Opisthobranchia:

Francisco J. Garcia Nudibranchia) from the Atlantic Ocean... 0 150

Jesus S. Troncoso

Frida Ben-Ami First report of the invasive freshwater snail Tarebia granifera (Lamarck,

1816) (Gastropoda: Thiaridae) from Israel 2... ee 156

NOOO? 1 peg Gabe bcs Gok Ble Cada EA: Gow hn, G GW GAGES hea, a hy gies aE Lays Aa WEE Lie Gow Wl Sha sae ar Shia 1s WY a dads 162

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THE NAUTILUS 120(4):119-130, 2006

age 119

Paryphantopsis (Gastropoda: Pulmonata: Charopidae) from the

Louisiade Archipelago of New Guinea

John Slapcinsky

Florida Museum of Natural History

University of Florida

Gainesville, FL 32611 USA

[email protected]

ABSTRACT

Recent surveys (2003-2004) of the terrestrial snail fauna of the

three largest islands in the Louisiade Archipelago, Misima,

Rossel and Sudest have uncovered a remarkable radiation of

Paryphantopsis, a diverse genus of charopid snails endemic to

New Guinea and nearby islands. Prev iously, only one species,

Paryphantopsis louisiadarum, here recognized as endemic to

Rossel Island, was known from the Louisiade Archipelago.

Three additional species were uncovered, all new and all ap-

pear to be endemic to single islands within the Louisiade Ar-

chipelago. The four species are described or redescribed using

information on shell, genital and radular anatomy. The terres-

trial mollusks of many of the other islands in the Louisiade

Archipelago have never been sampled and it is likely that di-

versity of Paryphantopsis and other land snails is underesti-

mated in the Louisiades.

Additional Keywords: Mollusca, mollusks, terrestrial, gastro-

pods, land snails, endemism, Papua New Guinea.

INTRODUCTION

This is the second in a series of reports on the results of

recent field surveys of terrestrial mollusks from Papua

New Guinea. The first report (Slapeinsky, 2005) de-

scribed six new species of the charopid genus Paryphan-

topsis from the eastern peninsula of mainland New

Guinea; this, the second, reviews Paryphantopsis species

collected during ten weeks of field surveys in January

2003 and April-May 2004 from the three lar gest islands

in the Louisiade Archipelago: Misimi 1 (St. Aignan), Ros-

sel (Yela), and Sudest (Vanatinai, Tagula). Paryphantop-

sis, a genus se alle snails endemic to New Guinea, is

comprised o: of twenty described species (Solem, 1970;

Slapcinsky, 2005) that are distributed from Western

a (Irian i a) to the Louisiade Archipelago. Only

one species, Paryphantopsis louisiadarum (Méllendortff,

1S99) was previously known from the Louisiade Archi-

pelago; it is the type of the genus Ilonesta (Iredale,

1941), later synonymized with Paryphantopsis (Solem,

1958).

The Louisiade Archipelago, a group of volcanic islands

and coral islets, with a total area of approximately 1600

kim”, lies about 300 km east of the New Guinea sete

and 400 km west of the Solomon Islands (Figure 1). The

archipelago is located on the southeastern extension of

the Owen Stanley Terrane, part of the East Papua Com-

posite Terrane ( ISPCT), a tectonic province composed of

at least four separate geological units with differing ages,

origins, and histories that appear to have assembled

northeast of modern New Guinea during the Paleocene,

62-57 Myr ago, and fused to the main body on the island

in the Late Oligoce ne to Early Miocene, 28-22 Myr ago.

The Louisiades are at least 15-20 million years old and

more likely were formed 40-60 million years ago and

have appi arently never had a land connection wath the

New Guinea mainland (Pigram and Davies, 1987). The

Louisiades physical isolation and great age combine to

provide considerable opportunity for the evolution of a

distinctive fauna. However, this fauna is poorly sampled,

especially for invertebrates, including terrestrial mol-

lusks. Only approximately 30 ia of land-snails are

known from the archipelago (Iredale, 1941), these were

collected during brief surveys in the mid to late 1g!"

Century. Nearly all of ns species appear to be re-

stricted to single islands, Low sampling intensity com-

bined with anticipated high levels of endemism suggests

that land snail diversity in the archipelago is under-

sampled.

MATERIALS AND METHODS

Specimens were hand-collected, drowned overnight, and

preserved in 75% ethanol. Gross anatomical dissections

were made in 75% ethanol using a dissecting microscope.

Radulae were isolated from dissected buccal masses us-

ing a saturated KOH solution. Scanning electron micro-

graphs of radulae were made using a Field Emission-

SEM. Line drawings of the genital anatomy were made

ein digital images, and measurements were taken using

an ocular micrometer. Shell and radular measurements

were made as figured in Slapcinsky (2005). Whorl count

was measured from the suture of the first whorl to the

body whorl and fractions of a whorl were determined

Page 120

THE NAUTILUS, Vol. 120, No. 4

3: 11°

E 153°

Figure lL.

Distribution of Paryphantopsis in the Louisiade

Papua

New Guinea

Louisiade

Archipelago

Sudest

E 154°

Archipelago, Papua New Guinea; L = P. louisiadarum, M = P.

misimensis, V = P. vanatinensis, Y = P. yelensis, 0 = other sites sampled .

with the aid of a cardboard circle divided into ten equal

parts of 36°. Spire diameter was the length of a straight

line passing from the apertural edge of the suture

through the middle of the apex to the opposite suture.

Diameter was the greatest width of the shell perpendicu-

lar to the Height was the greatest distance

between the apex and the base of the aperture measured

parallel to the shell axis. Spire height was measured from

the top of the body whorl to the apex of the shell. Ap-

erture width was the greatest distance from the columel-

lar edge to the outer edge of the aperture. Aperture

height was measured from the suture to the base of the

aperture, parallel to the shell axis. The following abbre-

viations are used in figures of genital anatomy: AT =

trium; DI = diverticulum; EP = epiphallus; OV = free

a. PE

pilasters: PR

thecal duct; SP

shell axis.

= penis; PG = prostate gland; PP = penial

SD = sperma-

and VD =

penial retractor muscle;

spermatheca; VA = vagina;

vas deferens. Terminology of vegetation types follows

Paijmans (1976). Specimens are de posited in the follow-

ing institutions: Bernice P. Bishop Museum, Honolulu

(BPBM); Florida Museum of Natural History, Gaines-

ville (UF); Natur-Museum Senckenberg, Frankfurt

(SMF); Papua New Guinea National Museum, Port Mo-

(PNGNM); Wroclaw University Museum of Natu-

(MNHW); Queensland Museum (QM).

resby

ral History

SYSTEMATICS

Family Charopidae Hutton, 1554

Genus Paryphantopsis Thiele, 1928

Type species: Flammulina (Paryphantopsis) lamelli-

gera Thiele, 1925, by original designation.

Paryphantopsis louisiadarum (Méllendortt, 1S99)

(Figures 2-9, Table 1)

John Slapcinsky, 2006 Page 12]

Qn9

Figures 2-9. Paryphantopsis louisiadarum. 2-4. Photographs of shell, UF 353425, diameter 10.6 mm. 5. Photograph of live

animal. 6-7. Camera lucida drawing of genitalia, UF 353426, maximum width 11.6 mm. 8-9. Scanning electron micrograph of radula,

UF 353426, field width of central and lateral teeth 61 xm, marginals 70 pm.

Paryphanta louisiadarum Mollendortf, 1899: 89; Méllendorff Description: The adult shell is depressed globose and

and Kobelt, 1902-1905: 17, pl. 3, figs. 1-3. large for the genus, 9.3-11.9 mm (mean = 10.3, see

se aac Mollendorff, 1899).—TIredale, 1941: Table 1 for sample size and standard deviation) in diam-

92-93. 7 ale. ee ae ey eA ei ade Sed ty

Paryphantopsis louisiadarum (MOllendorff, 1899).—Solem, eter and 6.7-9.0 a (mean = ; -O) in height, with 2.1-3.0

1958: 23: Solem, 1959: 156, pl. 12, figs. 10-11, pl. 13, fig (mean = 2.8) rapidly expanding whorls (Figures 2-4).

6: Solem, 1970: 259-260. The sutures are deeply impressed and the shell margin is

Page 122

THE NAUTILUS, Vol. 120, No. 4

evenly rounded. The spire is flat to slightly elevated 0.0—

0.2 mm (mean = 0.1). The body whorl descends slowly

near the aperture and the she I height/diameter ratio is

0.66-0.80 (mean = 0.73). The shell has 1.3 ev enly

rounded protoconch (nuclear) whorls, sculptured with

spiral rows of small pits, approximately 15 rows can be

seen on the apex of adult shells. These spiral pits con-

tinue on the teleoconch (post-nuclear) whorls eventually

merging into incised spiral striae that weaken towards

the aperture. The teleoconch whorls are also sculptured

with weak growth lines that are not accentuated by pe-

riostracal extensions; these are strongest on the body

whorl near the aperture. The protoconch and teleoconch

whorls are brown. The suture is darker brown and the

body whorl is usually irregularly maculated with darker

brown. The umbilicus is closed by a reflection of the

peristome. The aperture is large, compressed ovate, with

an aperture-width to aperture-height ratio of 0.91—0.98

(mean = 0.95).

The body color is bright yellow with green-black pig-

ment on the head and eyestalks exte nding i in two lateral

bands to the posterior of ‘the foot (Figure 5). These bands

are irregularly maculate anteriorly and ene ond ventrally,

often visible as spots on the sole of the foot. The yellow

fades to creamy-white in specimens preserved in ethanol.

The vas deferens remains narrow to the slightly swollen

head of the epiphallus (Figure 6). The epiphi allus does

not bear a diverticulum. The penis is 0.70 the length of

the gel and is apically robust, about 3 times the

width of the epiphallus at their junction, and tapers rap-

idly towards its base. Penis is sculptured with four pilas-

ters two of these are wide and two narrower. Each pi-

laster is regularly plicated perpendicular to the length of

the penis (Figure 7). The penial retractor muscle origi-

nates from the diaphragm and inserts on the basal 0.30 of

the epiphallus. The spermathecal duct is robust, narrow-

ing gradually from the basal 0.25 to the apical 0.25 and

remaining narrow until joining the ovate spermatheca.

The free oviduct joins the short vagina just above the

spermatheca.

The central teeth of the radula (Figure 8, center row)

are tricuspid, 12-13 xm wide and 15-16 jm long,

roughly the same shape as, but smaller than, the first

late mall teeth, which are 16-17 wm wide and 19-20 wm

long. The mesocones of both the centrals and first later-

als are blade shaped, apically robust, widest. slightly

Table 1

diameter, SH = spire height, SD = spire diameter, AH =

tude 6 mm. However,

Measurements in mm of undamaged adult shells of four species of Paryphantopsis, N = sample size, H =

aperture height, AD =

above their mid-point, and narrowing basally, joining the

rectangular basal plates close to, but not on, their poste-

rior edge. The mesocones of the central teeth do not

project beyond the anterior edge of the basal plates,

those of the lateral teeth project beyond the edge. The

ectocones are trigonal and short only 0.30 of the height of

the mesocones, joining the posterior edge of the basal

plates. The lateral teeth are barely asy mmetrical, their

endocones are only slightly taller than their ectocones.

The endocones of the ister ral teeth are slightly larger but

otherwise of similar shape to their ectocones. The first

ten teeth to the left and right of the central row are

similar to the first laterals, the next four teeth on either

side grade in shape and are difficult to classify as either

laterals or marginals. The last five are clearly marginal

teeth and are dorsoventrally compressed and tricuspid,

11-12 pm wide and 12-18 pm long (Figure 9). The

endocones of the marginal teeth are 0.70 to nearly the

same height as the mesocones and the ectocones vary

from less ean 0.50 to nearly 0.70 the height of the me-

socones. The endocones and mesocones of the marginal

teeth occasionally bear small notches or cusps near fhew

apices.

Lectotype: SMF 137274.

Paralectotype: SMF 165564 (1 specimen).

Remarks: Moéllendorffs description of Paryphanta

louisiadarum included a single set of measurements: ma-

jor diameter 9.5 mm, minor diameter 6.5 mim, and alti-

these measurements are difficult

to match to either of the two now slightly broken shells

of P. louisiadarum donated from his ecllection to Natur-

Museum Senckenberg, Frankfurt, and originally cata-

loged together as SMF 137274. One specimen has pat-

terns of missing periostracum similar to the shell figured

by Mollendorff and Kobelt (1902-1905). Labeling with

this shell indicates it was separated from the other speci-

men and marked “Lectotype” based on its similarity to

the shell figured by M6llendorff; later, Solem (1970) for-

mally published this lectotype designation. The other

specimen in the lot, now the paralectotype, was re-

catalogued as SMF 165564, Méllendorff's specimens of

Paryphanta louisiadarum came from Strubell, who also

height, D =

aperture width, W = number of whorls.

Species N H D SH SD AH AD W

P. lowisiadarum 50 Mean + SD 7.5+0.7 10.3 + 0.7 0.1401 3.6+0.3 6.2+0.4 6.6 40.5 28+0.]

Range 6.7-9.0 9.3-11.9 0.0-0.2 3.04.0 5.7-6.9 5.8-7.6 2.7-3.0

P. misimensis 1] Mean + SD 6.0 + 0.3 $.0 + 0.5 0.1+0.1 2.6 + 0.2 5.1 + 0.3 5.3 + 0.2 2.5 + 0.1

Range 5.7-6.6 7.2-8.7 0.0-0.3 93-3. 4.7-5.5 L.$—5.5 93-9.

P. vanatinensis 50 Mean + SD 5.8 + 0.5 7.9 +0.7 O.1 40.1 28+0.3 19+0.4 1.9 + 0.6 26+ 0.2

Range 5. 1-6.9 7.0-9.6 0:0-0:3 953.6 4.4—6.0 35.9-6.0 2.4-2.9

P. yelensis LO Mean + SD 2.7+0.3 3.9 + 0.3 O.1+0.1 1.6 + 0.2 1.9 +0.2 2.4+0.1 2.8 + 0.1

Range 9.4-3,] 3.64.5 0.0-0.2 1.5-1.9 1):6-2.1 2.3-2.6 2.7-3.0

John Slapcinsky, 2006

Page 123

sent specimens to Fulton. These specimens were distrib-

uted to other collections that eventually found their way

into museums, and are the source of UMMZ 127616 and

ANSP. 109257.

Type Locality: —Louisiaden (Louisiade Archipelago).

Other Material Examined: Papua New Guinea,

Milne Bay Province, Louisiade Islands, UMMZ 127616

(1 specimen): Rossel Island: ANSP 109257 (1 specimen);

UF 339009 (27 Sema base of Tachu G Gap, 569

meters lead 1.353° S, 154.223° E, J. Slapeinsky, 5

May 2004; UF 3 pcey 15 Getmaens PNGNM 005-001

(5 specimens), Lipuwopu at Lipu River, 320 meters al-

titude, 11.346° S, 154.221° E, ]. Slapcinsky, 12 May

2004; BPBM 268733 (5 specimens), MNHW MN 998 (5

specimens), UF 339011 (16 amas ns), UF 353425 (1

specimen), UF 353426 ( 1 specimen), ee Creek at

base of Tachu Gap, 63S meters altitude, 11.354° S$

154.223° E, } Slapcinsky, 6 May 2004; U scent (29

specimens), Tachu Gap below summit of Mount Rossel,

679 meters altitude, 11.356° S, 154.243° E, J. Slapcinsky,

4 May 2004; UF 339010 (4 specimens), W of former site

of G obubop Village, 255 meters altitude, 11.336° S,

154.221° E, J. Slapeinsky, 16 May 2004; UF 339013 (S

specimens), Wopu River upstream of trail crossing near

abandoned Yela Village, 280 meters altitude, 11.338° S,

154.224° E, J. Slapeinsky, 14 May 2004; SFM (5 speci-

mens), os MO76144 (5 specimens), UF 339014 (12

specimens), Yelebop Mountain S of Mount Rossel, 777

meters altitude, 11.357° S, 154.222° E, J. Slapcinsky, §

May 2004.

Habitat: Observed above 280 meters altitude in mixed

hill forest and mixed lower montane forest, active during

the day crawling on trees and shrubs from near ground

level to 2 m height.

Remarks: Within Paryphantopsis, P. louisiadarum is

similar only to P. globosa, P. misimensis, P. ubwamensis,

and P. vanatinensis in lacking all traces of periostracal

extensions on the growth lines. Paryphantopsis lousia-

darum differs from P. globosa in being smaller, having a

closed umbilicus and fewer whorls and differs from P.

misimensis and P. vanatinensis in having a penis that is

robust apically and narrow basally. It fccher differs in

body color pattern with two broad lateral bands not

sae) in P. misimensis that extend to the sole of the

foot unlike those of P. vanatinensis. The shell of P. loui-

siadarum is usually irregularly maculated with darker

pigment unlike the more uniformly colored P. misimen-

sis and P. vanatinensis. Paryphantopsis ubwamensis dif-

fers from P. louisiadarum in having a more tightly coiled

shell with a higher spire and an apical diverticulum on

the epiphallus.

Paryphantopsis misimensis new species

(Figures 10-17, Table 1)

Dene The adult shell is globose, large for the

genus, 7.2-8.7 mm (mean = 8.0, see Table | for sample

size and standard deviation) in diameter and 5.7-6.6 mm

mean = 6.0) in height, with 2.3-2.7 (mean = 2.5) rapidly

expanding whorls (Figures 10-12). The suture is deeply

impressed and the shell margin is evenly rounded. The

spire is flat to slightly elevate ad, 0.0-0.3 mm (mean = 0.1),

the body whorl descends slowly and regularly. Shell

he ight/diz meter ratio is 0.68-0.81 (mean = 0.75). There

are 1.3 evenly rounded protoconch whorls, sculptured

with spiral rows of small pits; approximately 13 rows can

be seen on the apex of adult shells. Rows of ‘spiral pits are

continued on the teleoconch, becoming weaker and

eventually merging to become incised spiral striae. Spiral

striae weaken be coming nearly obsolete on the final 0.25

of the body whorl where shell sculpture becomes pre-

dominated by weak and somewhat rounded growth lines

that do not bear periostracal extensions. The protoconch

and teleoconch whorls are uniformly brown, except for

some darker pigmentation in the suture. The umbilicus is

closed by a reflection of the peristome. The aperture is

large, ovate, with an aperture-width to aperture-height

ratio of O.S7—1.02 (mean = 0.97).

The body color is uniform yellow that fades to cream

in specimens preserved in ethanol; there are no lateral

bands on the foot or pigment on the eyestalks (Figure

13). The vas deferens narrows rapidly after the ple

gland and remains narrow until entering the slightly

swollen head of the epiphallus (Figure 14). The epiph ral-

lus is robust, especially basally where it is twice the di-

ameter of its mid-point, it does not bear a diverticulum,

and is 1.25 times penis length. The penial retractor

muscle originates on the diaphragm and is inserted near

the mid-point of the epiphallus. The penis is robust

throughout its length, approximately 0.25-0.30 larger

than che base of the epiphallus. Penis is sculptured

with two massive smooth pilasters (Figure 15). The

spermatheca is ovate, its duct is narrow throughout its

length expanding only slightly at the junction with the

free oviduct at the short vagina only slighily above the

atrium.

The central teeth of the radula (center row) are tri-

cuspid, 9-10 jzm wide and 16-17 wm long, smaller than

the first lateral teeth, which are 13-14 wm wide, 21-22

um long (Figure 16). The mesocones of the central teeth

do not project beyond the basal plate while those of the

lateral teeth do. The ectocones of the central and lateral

teeth are about 0.30 the height of the mesocones. The

laterals are tricuspid and slightly asymmetric with the

endocone of each lateral tooth slightly taller than the

ectocone. The endocones of the lateral teeth are slightly

larger but othenvise of similar shape to their ectocones

and both point towards their mesocones. The first seven

teeth to the left and right of the central row are similar

to the first lateral teeth, the next five teeth on either side

grade in shape and : 5 difficult to classify as either lateral

or marginal teeth. The last six are cl early marginal teeth

and are dorsoventrally compressed and tricuspid, with-

out accessory cusps, 11-12 xm wide and 15-15 tm long

(Figure 17). The endocones of the marginals are 0.70 the

height of the mesocones while the ectocones are shorter,

only 0.50 the height.

Page 124 THE NAUTILUS, Vol. 120, No. 4

—a_

Figures 10-17. Paryphantopsis misimensis. 10-12. Photographs of shell, Holotype UF 308234, diameter $.4 mm. 13. Photograph

of live animal. 14-15. camera lucida drawing of genitalia, UF 353422, maximum width 9.3 mm. 16-17. Scanning electron micro-

graph of radula, UF 353422, field width of central and lateral teeth 73 jum, marginal teeth 64 jum.

John Slapcinsky, 2006

Page 125

Holotype: UF 308234, F. Kraus,

Paratypes: BPBM 268734 (2 specimens), PNGNM

005-002 (5 specimens), UF 303579 (19 specimens), UF

303580 (7 specime iis), UF 353424 (1 specimen), UF

353422 (2 specimens), type locality, F. Kraus, 17 January

2003.

17 January 2003.

Type Locality: Papua New Guinea, Milne Bay Prov-

ince, Louisiade Archipelago, Misima Island, Oya Tau,

1014 meters altitude, 10.660° S, 152.629° E.

Habitat: Active during the day on trees and shrubs

within 2 meters of the ground in mixed lower montane

forest near the summit of Ova Tau.

Etymology: Named for Misima Island where this spe-

cies is presumed to be endemic.

Remarks: Sa onesie misimensis is similar only to

globosa, P. louisiadarum, P. ubwamensis, and P.

vanatinensis in lacking all traces of periostracal exten-

sions on the growth lines. Paryphantopsis misimensis dif-

fers from P. globosa in being smaller, having a closed

umbilicus and fewer whorls and differs from P. lowisia-

darum and P. vanatinensis in having a penis that is robust

throughout its length, not having dark lateral bands on

the foot, and having narrower radular teeth with en-

docones and ectocones that point towards the meso-

cones. Paryphantopsis ubwamensis difters from P. misi-

mensis in having a more tightly coiled shell with a higher

spire and an apical diverticulum on the epiphallus.

Paryphantopsis vanatinensis new species

(Figures 1S—25, Table 1)

Description: The adult shell is globose, larger than

average for the genus, 7.0-9.6 mm (mean = 7.9, see

Table 1 for sample size and standard deviation) in diam-

eter and 5.1-6.9 mm (mean = 5.8) in height, with 2.4-2.9

(mean = 2.6) rapidly expanding whorls (Figures 1S—20).

The suture is deeply impressed and the shell margin is

evenly rounded. The spire is flat to slightly elevated,

0.0-0.3 mm (mean = 0.1). Teleoconch “dbiorls descend

slowly and regularly until the final 0.70 of the body whorl

where it descends more ay. Shell height/diame ter

ratio is 0.77-0.91 (mean = 0.85). There are 1.3 evenly

rounded protoconch sable sculptured with spiral rows

of small pits; approximately 13 rows can be seen on the

apex of adult shells. These pits become larger and less

regular on the teleoconch whorls eventually fusing to

form incised spiral striae. Spiral striae w eaken slightly on

the final 0.25 of the body whorl where shell sculpture

becomes predominated by weak and somewhat rounded

growth lines that do not bear periostracal extensions.

Protoconch and teleoconch whorls are uniformly brown

except for some darker pigmentation near the aperture.

The umbilicus is closed by a reflection of the peristome.

The aperture is large, ovate, with an aperture-width to

aperture-height ratio of 0.92-1.12 (mean = 1.01).

The body color is bright yellow with green-black pig-

ment on the eyestalks and in two weak mid-lateral bands

on the anterior of the foot and two stronger mid-lateral

bands on the posterior half of the foot (Figure 21). These

bands are solid, not maculate, do not extend to the ven-

tral edge of the foot, and can not be seen on the sole of

the foot. In some individuals the anterior bands are lack-

ing. The yellow fades to creamy-white in specimens pre-

served in ethanol. The vas deferens narrows slightly to

the junction with slightly inflated cylindrical head of the

epiphallus (Figure 22). The epiphallus is approximately

0.50 the diameter of the penis, narrows only slightly ba-

sally and does not bear a diverticulum. The penial re-

tractor muscle is short, originating from the diaphragm,

and inserted at the basal 0.30 of the epiphallus. The

penis is about the same length as the epiphallus, and

sculptured with several rows of regularly plicate pilasters

(Figure 23). The atrium is moderate in size and of con-

sistent width. The spermathecal duct is robust remaining

wide for basal 0.50 then tapering gradually to the junc-

tion with the ovate spermatheca. The free ein is

about 0.50 the diameter of the spermathecal duct where

they join. The vagina is of moderate length about the

same length as the atrium.

The conte teeth of the radula (middle row) are sym-

metrically tricuspid, 13-14 jum wide and 17-18 jum long.

The slightly asymmetrical lateral teeth are otherwise

similar in shape to the central teeth but are slightly

wider, 15-16 wim wide, and nearly identical in length,

17-18 um long (Figure 24). The bluntly conical ani

erect mesocones of the central and lateral rows join their

basal plates near the posterior edge and project to or

slightly beyond the anterior of their basal plates. The

ectocones of both the central and lateral rows are trigo-

nal and short, about 0.50 of the height of the mesocones,

and join the posterior edge of their basal plates. The

endocones of the lateral tooth are slightly larger but oth-

erwise of similar shape to their ectocones. The first ten

teeth to the left and right of the central row are clearly

lateral teeth; the next five on either side grade in shape

and are difficult to classify as either laterals or marginal

teeth. The last five are clearly marginal teeth and are

dorsoventrally compressed, tricuspid, about 11-14 pm

wide and 13-19 wm long (Figure 25). The endocones are

a bit more than 0.70 the heights of the mesocones while

the ectocones are slightly shorter, about 0.70 the height

of the mesocones.

Holotype: UF 353421, J. Slapcinsky, 19 April 2004.

Paratypes: Papua New Guinea, Milne Bay Province,

Louisiade Archipelago, Sudest Island (Vanatinai,

Tagula): BPBM 268735 (10 specimens), MNHW MP

989 (5 specimens), PNGNM 005-003 (10 specimens),

SFM (10 specimens), QM MO76145 (5 a oennnnies UF

339019 (68 specimens), UP 547729 (3 specimens), UF

393420 (1 specimen), UF 353423 (1 specimen), type lo-

cality, J. Slapcinsky, 19 April 2004; UF 339018 (41 speci-

mens), base of Mount Boa Mount Rio) on the Esiraba

River, 120 meters altitude, 11.492° S, 153.413° E, J.

Slapcinsky, 15 April 2004; UF 339017 (37 specimens),

near Avarumolo Rock Shelter, 150 meters altitude,

11.490° S, 153.420° E, J. Slapcinsky, 23 April 2004.

Page 126 THE NAUTILUS, Vol. 120, No. 4

Figures 18-25. 9 Paryphantopsis vanatinensis. 18-20. Photographs of shell, Holotype UF 353421, diameter 7.6 mm. 21. Photo-

graph of live animal. 22-23. Camera lucida drawing of genitalia, UF 353423, maximum width 8.9 mm, 24-25. Scanning electron

micrograph of radula, UF 353423, field width of central and lateral teeth S7 zm, marginals 52 jm.

Type Locality: Papua New Guinea, Milne Bay Prov- Habitat: Found active during the day on trees and

ince, Louisiade Archipelago, Sudest Island (Vanatinai, shrubs from near ground level to 2 meters in mixed hill

Tagula), Emua Peak, just W of the summit of Mount Riu forest and mixed lower montane forest from 120 meters

(Mount Rio), 725 meters altitude, 11.507° S, 153.431° E. to 725 meters altitude.

John Slapcinsky, 2006

Page 127

Etymology: Named for Vanatinai Island where this

species is presumed to be endemic.

Remarks: = Paryphantopsis vanatinensis is similar only

to P. globosa, P. louisiadarum, P. misimensis, and P. ub-

wamensis in lacking all traces of periostracal extensions

on the growth lines. Paryphantopsis vanatinensis differs

from P. globosa in being smaller, having a closed umbi-

licus and fewer whorls and differs from P. louisiadarum

and P. misimensis in having a penis that is relatively nar-

row throughout its length and havi ing dark lateral bands

that are not maculate and do not exte nd to the sole of the

foot. Paryphantopsis ubwamensis differs from P.

vanatinensis in having a more tightly coiled shell with a

higher spire and an apical div erticulum on the epiphal-

lus.

Paryphantopsis yelensis new species

(Figures 26-32, Table 1)

Description: The adult shell is globose to depressed

globose, small for the genus, 3.6-4.5 mm (mean = 3.9,

see Table 1 for sample size and standard deviation) in

diameter and 2.4-3.1 mm (mean = 2.7) in height, with

2.7-3.0 (mean = 2.7) rapidly expanding whorls (Figures

26-28). The shell is wider and slightly angular below the

mid- point and its suture is deep, sometimes appenine

nearly adnate. The spire is flat to elevated, 0.0-0.2 mm

(mean = 0.1). Teleoconch whorls descend regularly and

shell height/diameter ratio is 0.62-0.73 (mean = 0.68).

There are 1.2 rounded protoconch whorls; approximately

13 rows of spiral lirae can be seen on the apex of adult

shells. These lirae are crossed by stronger sharp axial

lines forming a lattice pattern. The teleocotich is sculp-

tured with spiral rows of pits that become less regular

malleations on the final third of the body whorl. This

shell sculpture is somewhat obscured by periostracal ex-

tensions approximately every 6-7 growth lines that are

accentuated with processes at the angled shell margin.

These processes are often worn and can be missing from

some adult shells. The protoconch and teleoconch whorls

are uniformly brown. A reflection of the peristome closes

or nearly closes the umbilicus. The aperture is large,

nearly circular except at the angled periphery and has an

aperture-width to aperture- height ratio of 0.69-0.93

(mean = 0.80).

In life the body color is bright yellow, there are no

lateral patches on ‘the foot, the ey estalks are dark black-

brown, the yellow fades to cream in specimens preserved

in ethanol. The vas deferens narrows toward the j junction

with the inflated spherical head of the epiphallus. Im-

mediately below the head of the epiphallus there is a

finger shaped apical diverticulum that is roughly 2 the

length of and nearly the same diameter as the e piphe illus

(Figure 29). The epiphallus is approximately the same

length h as the penis. The penial retractor muscle is long,

originating from the diaphragm, and inserted just be low

fhe: mid point of the epiphallus. The epiphallus is roughly

half the diameter of the penis. The penis is widest 0.30 of

the way down from the apex and narrows slightly towards

the apex and towards the base where it is 0.70 the diam-

eter of the atrium at their junction. Penis sculptured with

a narrow pilaster that widens basally (Figure 30). The

atrium is average to short, about 0.50 the le ngth of the

relatively long vagina. The spermathecal duct, free ovi-

duct, and vagina are all of similar diameter at their junc-

tion. The spermathecal duct is relatively narrow basally

and tapers slowly and evenly until joining the spherical

spermatheca.

The central teeth of the radula (center row) are tri-

cuspid, 7-8 fxm wide and 12-13 pm long, of similar

shape and length but slightly narrower than the first lat-

eral teeth, 8-9 wm wide and 12-13 jum long ( Figure 31).

The mesocones of both the central and first late val teeth

are relatively short and blunt, not tapering until the

rounded tip of the cusp and not extending beyond their

basal plates. They appear particularly shit because they

project nearly perpendicularly from their basal plates

and not as much anteriorly. Mesocones of the central and

lateral teeth are attached toward the middle of their basal

plates. The ectocones of the central te eth and the nearly

symmetric ectocones and endocones of the laterals are

trigonal and about 0.50 the height of the mesocones. The

eadocinies of the lateral teeth are slightly larger but oth-

erwise of similar shape to their ectocones. The { first five

teeth to the left and right of the central row are similar

to the first lateral teeth, the next three on either side

grade in shape and are difficult to classify as either lat-

els or marginal teeth. The last five are clearly marginal

teeth and are dorsoventrally compressed and tricuspid,

7-9 wm wide and 7-8 wm long (Figure 32). The ecto-

cones of the marginal teeth are only slightly shorter than

their endocones, which are only ‘slightly shorter than

their mesocones.

Holotype: UF 353427, J. Slapcinsky, 12 May 2004.

Paratypes: Papua New Guinea, Milne Bay Province,

Louisiade Archipelago, Yela Island (Rossel Island): UF

339016 (15 specimens), UF 353428 (1 specimen), type

locality, J. Slapcinsky, 12 May 2004; UF 339120 (1 speci-

men), Wopu River upstream of trail crossing near aban-

doned Yela Village, 280 meters altitude, 11.338° S,

154,224° E, J. Slapcinsky, 14 May 2004; UF 339119 (1

specimen), former site of Copan Village, 275 me ters

altitude, 11.335° S, 154.222° E, J. a peinsky, 1 3 May

2004; UF 342911 (2 specimens). \ ’ of former site of

Gopubop Village, 255 meters suede 11.336° S,

154.221° E, J. Slapcinsky, 16 May 2004.

Type Locality: Papua New Guinea, Milne Bay Prov-

ince, Louisiade Archipelago, Yela Island (Rossel Isl: ind),

Lipuwopu at Lipu River, 320 meters altitude, 11.546° S

154.221° E.

Habitat: All specimens were collected in mixed

hill forest from 275 to 285 m altitude. Individuals

were observed active during the day on boulders that

were densely encrusted with algae, mosses and lichens,

and, less commonly, on dead logs and twigs near the

sround.

Page 128

THE NAUTILUS, Vol. 120, No. 4

Figures 26-32.

lucida drawing of genitalia, UF 353428, maximum width 4.7 mm

Remarks: = Paryphantopsis yelensis differs from. all

other Paryphantopsis from the Louisiades, P. lowisia-

darum, P. misimensis, and P. vanatinensis, and from the

lowland species from the mainland, P. lebasii and P.

yawii, in having an apical diverticulum on the epiphallus.

Paryphantopsis yelensis. 26-28. Photographs of shell, Holotype UF 353427, diameter 4.3 mm. 29-30. Camera

2 QQ

. 31-32. Scanning electron micrograph of radula, UF 353428, field

width of central and lateral teeth 47 2m, marginal teeth 27 wm.

Paryphantopsis yelensis differs from all other species of

Paryphantopsis other than P. globosa and P. striata in

having protoconch sculpture of axial and spiral lirae

rather than spiral pits. However, P. globosa and P. striata

have roughly equal axial and spiral lirae while P. yelensis

John Slapcinsky, 2006

Page 129

has stronger axial sculpture. Of the species for which the

radular morphology is known, the origin of the meso-

cones from the center of the basal plate in the central

and lateral teeth of P. yelensis is similar only to P. lebasii

and P. yawii.

Etymology: Named for Yela Island where this species

is presume sd to be endemic.

DISCUSSION AND CONCLUSIONS

New Guinea lies at the leading edge of the Australian

plate and is geologically complex, with much of the east-

ern and northern edge of the island and nearby satellite

islands being composed of accreted terranes ( (Pigr am and

Davies, 1987). This complex geology has promoted the

development of a unique and diverse biota that is still

being discovered. Terrestrial invertebrates including

snails are especially poorly known, being both ade

equately sampled and diverse as evidenced by recent

colle cting in eastern Papua New Guinea. Rece nt collect-

ing has uncovered high diversity and endemism in the

genus Paryphantopsis, with all isolated mountains

sampled having eve suites of endemic and previously

undescribed species (Slapcinsky, 2005).

The islands off New Guinea including the Louisiades

are also rich in endemic Paryj shantopsis species, al-

though only one of these, P. louisiadarum, was previously

described (Méllendorff, 1899) and later localized to Ros-

sel Island (Solem, 1958) where it appears to be endemic.

The Louisiades are also very poorly sampled with only

approximately 30 species of terrestrial snails previously

known from the archipelago (Iredale, 1941). The ongo-

ing brief surveys of a smal portion of the three large st

islands in the Louisiade Archipelago have increased aihe

known diversity of Paryphantopsis in the Louisiades by

300%. In addition to Paryphantopsis, species from sev-

eral other charopid genera and other families were also

collected; these will be treated in later publications. The

family Chi mie ve, previously considered to be a minor

component of the terrestrial mollusk fauna of New

Guinea, with relatively few species and genera (Solem,

1983), may be among ‘the most diverse familie s on New

Guinea and its sate ‘lite islands, rivaling the spectacular

radiations exhibited by this family in the oceanic Pacific

(Solem, 1983).

Two groups of Paryphantopsis appear to have colo-

nized the Louisiade Archipelago. One group, P. louisia-

darum, P. misimensis, and P. vanatinensis share similar

large, globose shells that have nearly flat spires and large

apertures, are sculptured with spiral rows of pits that fuse

to form incised spiral striae, and do not bear periostracal

extensions or processes on the growth lines. A second

group includes only P. yelensis a small species with

strong axial sculpture on the protoconch and periostracal

extensions with marginal processes on the growth lines of

the teleoconch and with an apical diverticulum on the

epiphallus. The relationship of both Louisiade clades

with species from mainland Papua New Guinea is not

clear. The three large species from the Louisiades, P.

louisiadarum, P. misimensis, and P. vanatinensis lack pe-

riostracal extensions on the growth lines similar only to P.

globosa and P. ubwamensis from mainland Papua New

Guinea; however, they also lack an apical diverticulum

similar only to P. lebasii and P. yawii from the adjacent

Papuan Peninsula. The relationship of P. yelensis to

mainland species of Paryphantopsis is equally cryptic.

The radula in this species has central and lateral teeth

with mesocones that originate from near the center of

the basal plates, a trait that is thus far known only from

P. lebasii and P. yawii, however unlike those species, P.

yelensis possesses an apical diverticulum. Although ana-

tomic evidence suggests two origins for Paryphantopsis

in the Louisiade Archipelago, additional anatomic and/or

genetic characters would be necessary to determine the

relationships of these species to those from mainland

New Guinea.

ACKNOWLEDGMENTS

I thank the landowners of Araetha, Tcheme, Damemu,

and Liak for permission to work on their land and for

field assistance; F. Kraus, F. Malesa, D. Mitchell, and G.

Shea for additional field assistance; G. Kula and D.

Mitchell of Conservation International for providing lo-

gistical support and advice; sae New Guinea National

Museum and Art Galle ry for providing in-country col-

laborative assistance; Papui t New Guinea Department of

Environment and Conservation, Papua New Guinea Na-

tional Research Institute, and Milne Bay Provincial Gov-

ernment for permission to work in Milne Bay Province;

K. Kelley, Electron Microscopy Core I ees Uni-

versity of Florida for ime ging radulae; Neubert for

photographing labels and type material of Paryphantop-

sis louisiadarum; L. Appleton, P. Callomon, R. Jenssen,

D. Potter, and M. She -a for facilitating my study of speci-

mens in their care; and J. Worsfold for sharing biblio-

graphic information. Fieldwork for this research was sup-

ported by National Science Foundation grant DEB

0103794 a the University of Florida Foundation.

MeGinty Endowment. F. Kraus, T. Pearce. F. G.

Thompson, and an anonymous reviewer suggested im-

provements to earlier drafts of this manusc ript.

LITERATURE CITED

Iredale, T. 1941. A basic list of the land Mollusca of Papua. The

Australian Zoologist 10: 51-94.

Mollendorff, O. 1899. Neue arten aus der Strubell’schen

sammlung. Nachrichtsblatt der Deutschen Malakozoolo-

sischen Gesellschaft 31(5): S9-92

Mollendorff, O. and W. Kobelt, 1902-1905. Die raublungen-

schnecken (Agnatha). Erste abtheilung: Rhytididae et En-

neidae. Syste smatisches Conchylien-Cabinet von Martini

und Chemnitz I, 12, B: 1-362; plts 1-41

Page 130

THE NAUTILUS, Vol. 120, No. 4

Paijmans, K. 1976. Vegetation. In: Paijmans, k. (ed.) New

Guinea Vegetation, Australia National University Press,

Canberra, 212 pp.

Pigram, C. J. and Davies, H. L. 1987. Terranes and the accre-

tion history of the New Guinea orogen. BMR Journal of

Australian Geology and Geophysics 10: 193-211.

Slapeinsky, J. 2005. Six new species of Paryphantopsis (Gas-

tropoda: Pulmonata: Charopidae) from the Papuan Pen-

insula of New Guinea. The Nautilus 119: 27-42.

Solem, A. G. 1958. Endodontide landschnecken von Indone-

sien und Neu Guinea. Archiv fiir Molluskenkunde 87; 19—

26.

Solem, A. G. 1959. On the family position of some Palau, New

Guinea, and Queensland land snails. Archiv fiir Mol-

luskenkunde 88: 151-158; pls. 12, 13, 2 figs.

Solem, A. G. 1970. The Endodontid land snail genera Pilsbry-

charopa and Paryphantopsis (Mollusca: Pulmonata). The

Veliger 12; 239-264.

Solem, A. G. 1983. Endodontoid land snails from Pacific Is-

lands (Mollusca: Pulmonata: Sigmurethra) Part II, Fami-

lies Punctidae and Charopidae, Zoogeography. Field Mu-

seum of Natural History, Chicago, ix + 336 p.

Thiele, J. 1925. Mollusken vom Bismark-Archipel, von Neu-

Guinea und Nachbar-Inseln. Zoologische Jahrbiicher 55:

119-146.

THE NAUTILUS 120(4):131-138, 2006

Page 13]

Daedalochila lithica and Daedalochila dorfeuilliana (Gastropoda:

Polygyridae) in Arkansas, USA: morphology, distribution,

and habitat

Brian F. Coles

Mollusca Section

Gerald E. Walsh

3065 N. Dorchester Drive

Fayetteville, AR 72703 USA

Department of Biodiversity

National Museum of Wales

Cathays Park, Cardiff, CF 10 3NP

UNITED KINGDOM

ABSTRACT

Shell and genital morphology, geographic distribution, and

habitat of two polvgyrid species, Daedalochila lithica and Dae-

dalochila dorfeuilliana in Arkansas are described. The two spe-

cies have been confused because their shells are similar and the

anatomy of D. lithica has not been previously described. In this

paper, the genital anatomy of D. lithica is described and shell

and genital anatomy of the two species are compared. No sta-

tistically significant differences were found in 15. shell-

morphometric characters. The only constant difference in shell

morphology was in the es palatal lamella of D. lithica, which

is less immersed than that of D. dorfeuilliana. Anatomically, the

species differed in diameter of the penis, development of the

epiphallus, thickness of the vas deferens at its junction with the

prostate gland and ratio of vagina length to diameter. Genital

anatomy indicates that both species belong to the Daedalochila

plicata group. They are found in oak-hickory forest: D. lithica

in areas where vegetation has been cut; D. dorfeuilliana in

undisturbed old growth forest. These data support the view that

D. lithica is a taxon distinct from D. dorfeuilliana and other

polygyrid species.

INTRODUCTION

Daedalochila dorfe uilliana (I. Lea, 1838) is a widely dis-

tributed land snail of the Ozark and Ouachita Mountain

regions of Arkansas, Oklahoma, and Missouri, extending

into Louisiana, Texas, Kansas and Illinois (Pilsbry, 1903;

Pilsbry and Ferriss, 1906; Hubricht, 1985). The species is

placed in the Polygyra ea group of the Polygyridae,

Polygyrini (Pilsbry, 1940), and is distinguisl red from

other members of the group by its geographical range,

and by the palatal and basal lamellae of ap-

proximately equal size (Pilsbry, 1940). Until Hubricht’s

description of Daedalochila lithica (enache 1961),

variation of shell morphology within this diagnosis had

been accepted as falling within the range of variation of

D. dorfeuilliana (Branson, 1970). The distribution of D

lithica, extending from north central and west central

Arkansas into southeastern Oklahoma (Hubricht, 1985),

falls entirely within the range of D. wells uilliana. Hu-

bricht (1961) described D. lithica as related to D. dor-

feuilliana but differing in details of the apertural lamel-

lae: “Polygyra lithica is related to Polygyra dorfeuilliana,

differing in the teeth. The parietal tooth is lower and is

veuinded rather than squarish. The lip teeth are smaller

and are not deeply immersed.”

Hubricht (1985) described D. lithica and D. dorfeuil-

liana as “calciphiles found under logs, rocks, and leaf

litter in dry upland woods” but they are never found

together.

Hubricht (1961) did not report the anatomy of D.

lithica. However, Pratt (1981b) placed the species in the

genus Millerelix Pratt, 1981 “because of its position as a

precise intermediate between Plolygyra| mooreana and

Plolygyra| dorfeuilliana in shell characters”; both Dae-

dalocbils mooreana (Binney, 1857) and D. dorfeuilliana

having been assigned to Millerelix on the basis of shell

characte ars ancl geniti ul anatomy (Pratt, 198la, b). Subse-

quently, Emberton (1995) pli iced all members of the

Polyg yr plicata group into Millerelix, separating the

species of Millerelix sensu Pratt, 19S1b, from other mem-

bers of the genus by erection of the subgenus Prattelix

Emberton, 1995. Coles and Walsh ( 2006) dissected

the well-defined species of the Polygyra plicata group

and expressed doubt as to the validity of placing all mem-

bers of the group in Millerelix because of the inconsis-

tency of diagnostic generic features. Consequently, the

species were referred to the senior genus Dae -dalochila

(Beck, 1837), However, D. lithica was not dissected by

Coles and Walsh (2006) because wet-preserved type ma-

terial did not exist and because of the uncertainty of

identification of new material as noted above. The

dissected specimen of D. dorfeuilliana illustrated in

Coles and Walsh (2006) came from outside the range

of D. lithica as given by Hubricht (1985). We were un-

able to examine material from the type locality of D

dorfeuilliana because that locality is unknown (Pilsbry,

1940). ,

Page 132

THE NAUTILUS, Vol. 120, No. 4

We have used type material of D. lithica plus material

identified by Hubricht as D. dorfeuilliana and D. lithica

to examine Hubricht’s (1961) concept of the two species

in more detail. Live material was obtained for study of

the anatomy and habitat preferences so as to identify

which characters of the shell and anatomy can be used to

define D. dorfeuilliana and D. lithica and to clarify the

relationship of the two species within the Daedalochila

plicata group.

MATERIALS AND METHODS

Material Examined:

four adult (based on development of the aperture and

apertural lamellae) paratype shells from the University of

Michigan Museum of Zoology (UNMZ), Ann Arbor,

Catalog Number UNMZ 205895; 19 lots (291 specimens

of adult shells) collected between 10 April 1936 and 1975

(no day/month given) from the Hubricht Collection at

the Field Museum of Natural History (FMNH), Chi-

cago, Catalog Numbers FMNIL 25 56348256362 and

FMNH 256369-256370; 14 lots (132 specimens of adult

shells) collected by the senior author between 10 April

1996 and 14 Septe smber 2000 from seven Arkansas coun-

ties. Daedalochila dorfeuilliana: 50 lots (299 adult shells)

from the Hubricht Collection of the FMNH, Catalog

Numbers FMNH 255102-255148 and FMNH 256377;

96 lots (142 adult shells) collected by the senior author

between 12 June 1995 and 24 March 2000 from nine

Arkansas counties.

Shell Measurements: Shell characters that were mea-

sured are shown in Figures 1-4, except body whorl

height, which was measured immedi ately behind the

outer lip of the aperture. Measurements were made to

the nearest 0.01 mm with an electronic digital caliper,

accuracy of + 0.03 mm. Ratios of shell height/diameter,

number of whorls/shell diameter, inside diameter of ap-

erture/outside diameter of aperture, width of parietal

tooth/height of parietal tooth and diameter of umbilicus/

diameter of shell were calculated. Student’s two-sample

t test was conducted for each shell variable and box plots

graphed on a Texas Instruments TI-S4 Plus graphing

calculator coupled with a Power Macintosh G3 desktop

computer.

Genital Anatomy: Specimens were drowned over-

night in sealed containers of degassed (boiled and

cooled) water and preserved in 70% ethanol. Dissections

were performed at 20-40x magnification. The shell was

removed from the body, the animal opened from near

the genital pore to the lung, and the tissues teased apart

while submerged in 70% ethanol. The right ocular re-

tractor muscle was cut and partially removed to aid view-

ing the basal penis. The penis was opened with a sharp-

ened spear-point needle from near its junction with the

atrium/vagina to the attachment of the penial retractor

muscle. Sections of the penis were obtained by cutting

the opened penial tubes where indicated in the figures

and allowing the remaining tissues to reform to their

Daedalochila lithica: one lot of

Shell dimensions measured. 1. d, diameter of

shell; 2. du, diameter of umbilicus; 3. h, height of shell; 4. od,

outside diameter of aperture; id, inside diameter of aperture;

wp, width of parietal lamella; hp, height of parietal lamella.

Daedalochila dorfeuilliana, FMNH 308106, 8.4 mm diameter.

Figures 1-4.

approximate original cylindrical forms. Drawings were

made by eye, taking particular care to ensure that relat e

aa catrgad of the organs were accurately reproduced,

and with reference to an appropriate scale.

Dissected Specimens: Daedalochila lithica (Figures

5-16), eight specimens from the following Arkansas lo-

calities: Boone County, Buffalo National sa Rush

Mountain Trail, 29 April 2004, G. Walsh, UF 376769,

two specimens dissected and measured ( pen ee

14); Boone County, Stonington Road, NE of New Hope,

2.1 ee N of Highway 14, 22 September 2004, G.,

Walsh, UF 376765, one specimen dissected and mea-

sured (Figure 16); Carroll County, Lake Leatherwood

Park, | 3 April 2004, G. Walsh, UF 376816, two speci-

mens eel one measured (Figures 12, 13); Madison

County, W Lae oo State Park, 22 September

2004, G. Walsh, UF 376766, two specimens dissected

and measured ( ee s S—10, 15); Marion County, Buf-

falo National River at Buffalo Point, B. Coles, 20 January

2003, UF 376767, one specimen dissected and measure od

(Figure 11), Daedalochila dorfeuilliana (Figures 17- 24),

seven specimens from the following Arkansas localities

were dissected and measured: Hempstead County, Lake

Millwood at Saratoga Landing, B. Coles, 5 January

2003, FMNH 293240, one specimen (Figures 17, 1S)

and UF 376768, one specimen (Figures 19, 20);

Faulkner County, River Plantation Golf Course (High-

way 365), Morgan, B. Coles, 10 October 2005,

NMW.Z.2005.011.03295, one specimen measured and

illustrated (Figures 21, 22); NMW.Z.2005.011.03296,

G. E. Walsh and B. F. Coles, 2006

Page 133

two specimens, Faulkner County, as above (dissected but

not illustrated); Carroll County, pa aver Dam Overlook,

15 and 20 March 2005, G. Walsh, UF 376770, two speci-

mens (Figures 23, 24),

Distribution and Habitat: Geographical distribu-

tions of D. lithica and D. dorfeuilliana in Arkansas were

compiled from the Hubricht collection at the FMNH

and the authors’ collections. Environmental conditions

were noted for each collection site visited by the authors.

Disposition of Specimens: — All shells of the senior au-

thor have been deposited in the FMNH. These include

7 lots of D. lithica (FMNH 308144—30S8150) and 40 lots

of D. dorfeuilliana (FMNH 308104—308143). Collections

of the junior author have been deposited in the Florida

Museum of Natural History (UF), Gainesville, Florida,

and the National Museum of Wales (NMW), Wales:

catalog numbers are presently available only for critical

dissected material given below,

RESULTS

Shell Characters: Visual examination of Hubricht’s

material of D. lithica and D. dorfeuilliana in the FMNH

confirmed the general features of his description of D.

lithica and its comparison with D. dorfeuilliana (Hu-

bricht, 1961). Thus, shells of the two species are similar

in apical and umbilical views (Figures 25-28), and the

structure of the apertural lamellae of many examples of

D. lithica (e.g., Figure 29) shows the low, rounded pari-

etal lamella and conical “not deeply immersed” apertural

lip lamellae that conform to Hubricht’s (1961) descrip-

tion of the species. However, detailed examination of the

several hundred shells of D. lithica and D. dorfeuilliana

that comprise the material identified by Hubricht

showed that there was considerable variation in the form

of the apertural lamellae. For example, a higher, less

rounded parietal lamella was often present in specimens

identified as D. lithica (Figure 30), the shape being simi-

lar to the “squarish” parietal lamella of D. dorfeuilliana

(Figure 31) as described by Pilsbry (1940). Conversely,

the parietal lamella of D. dorfeuilliana (Figure 32) was

often similar to that of D. lithica. In alditon, the basal

lamellae of both species were similar, i.e., conical and not

deeply placed within the aperture (Figures 29-32), and

did not appear to offer reliable diagnostic characters.

Thus, of Hubricht’s description of D. lithica, the depth of

immersion of the palatal lamella appears to provide the

only consistent shell character for separation of the two

species (Figures 29-32). The palatal lamella of D. lithica

is always placed more pee ly on the - whereas

hat of D. dorfeuilliana is always well below the lip.

Reexamination of Hubricht’s material confirme A that

specific assignments could be made on this basis. Simi-

arly, shells in the collection of the authors showed simi-

ar variation of form of the apertural lamellae, but all

including wet preserved material) could be assigned to

he palatal lamella.

D. lithica or D. dorfeuilliana on the basis of the de pth of

The results of a more formal analysis of variation of

shell parameters are shown in Figures 33-47, and the

Arkansas counties from which specimens were taken for

this analysis are shown in Figures 4549. Both species

showed considerable variation in all shell parameters

measured, with overlap of the ranges of the values, and

no statistical differences were found (P>+0.05).

Genital Anatomy: = Daedalochila lithica: Atrium short,

approximately twice the diameter of the basal penis (Fig-

ures 5, 12, 14-16); vagina of similar length, as broad in

diameter as the basal penis (Figures 14, 16) or slightly

inflated (Figures 5, 12, 15), its length approximately

equal to its diameter (Figure 16); free oviduct approxi-

mately half the length of the penis (Figures 5, 12, 14-16);

penis elongate, tape ring to a weakly defined, variously

flexed (Figures 5, 14-16) or coiled (Figure 12) epiphal-

lus, without appendix or flagellum; le nngth 2.63-3.89 min

(mean = 3.44 mm+0.53 mm, n = 7) including the

epiphallus; maximum width at mid-length, but basal pe-

nis only slightly narrower; widtlength of penis 0.07—

0.18 (mean = 0.13+0.04, n = 7 ): pe nial retractor muscle

terminal, variable in length a form; internally, the pe-

nis bears two fleshy pilasters that become weak in the

terminal penis/epiphallus (Figures 7, 9-11); in the ter-

minal 1/10 of the penis there is a minute papilla approxi-

mately 0.05 mm long attached to the penial walls and

pilaste rs by thickenings of the tissues (Figures LO, 11);

presence ofa papilla was not confirmed for all dissecte .d

specimens; vas deferens narrow, expanding from ap-

proximately mid length to reach its maximum diameter

(approximately twice its minimum diameter) proximally,

at the junction with the prostate gland (Figures 6, 13): no

other papillae, glandular regions, ealeed: regions or ad-

ditional features were asible in the penis at 40x magni-

fication.

The genital anatomy of D. dorfeuilliana has been de-

scribed ond illustrated for specimens from Tulsa County,

OKahoma, Pratt (19S1a, reproduced in Emberton, 1995)

and Hempstead County Arkansas (Coles and Walsh,

2006). Dissection of additional material is given in Fig-

ures 19-24, including a copra n from within the range

of D. lithica ( Figures 93, 24). These dissections confirm

the features of the genital iene of D. dorfeuilliana as

described by us (Coles and Walsh, 2006): notably the

penis widest in mid length where the pilasters are well-

developed, with a narrow base, pe nis length (including

epiphallus) 2.15-3.77 mm (mean 2.82 + 0.68 mm, n = 4),

width/length 0.13-0.15 (mean 0.15 + 0.01, n = 4); and

the proximal vas deferens at the junction with the pros-

tate gland greatly enlarged, approximately fourfold its

minimum diameter (Figures 21, 23). In ‘addition, the

length of the vagina is approximately twice its diameter

(Figure 21), and a minute papilla approximately 0.05 mm

long was found in the terminal 1/10 of the penis of a

specimen from Lake Milwood (Figures 19, 20), although

this was not confirmed for other specimens. The wvell-

differentiated and strongly reflected terminal penis (re-

ferred to here as the epiphé ulus, Figures 17, 23, 24) was

Page 134 THE NAUTILUS, Vol. 120, No. 4

G. E. Walsh and B. F. Coles, 2006 Page 135

Figures 5-24. Genital anatomy of Daedalochila lithica (Figures 5-16) and Daedalochila lithica (Figures 17-24) from Arkansas. 5.

D. lithica Boone ( Jounty, 29 April 2004, UF 376769. 6. Same, detail of junction of vas deferens (vd) and prostate gland (pt). 7. Same,

penis opened to show the two pilasters and the papilla (pa) in the terminal penis. 8. D. lithica, Madison County, 22 September 2004

UF 376766, showing a stout penis (p) and a short penial retractor muscle (pr). 9. Same, detail of penis opened to near the penial

retractor muscle. 10. Same, apical penis opened to the penial retractor muscle, showing obsolete apical pilasters and detail of terminal

penial pi apilla. LL. D. lithica, Marion County, 20 January 2003, UF 3876767, apical penis, opened, showing obsolete apical pilasters

and de = il of terminal penial papilla. 12. D. lithica, Car roll County, 3 April 2004, UF 376816, distal genitalia showing narrow, coiled

penis (p) and sections of penis as indicated in A and B. 13. D. lithic ‘a, Carroll County, 3 April 2004, UF 376816 (second specimen)

detail of junction of vas deferens and prostate gland. 14-16. D. lithica, distal genitalia Senay. variation of penis. 14. From Boone

County, 29 April 2004, UF 376769 iserone specimen). 15. From Madison County, 22 September 2004, UF 376766 (second

specimen). 16. From Boone County, 22 Septe ne 2004, UF racaon 17. D. dorfeuilliana, Hempstead County, 5 January 2003,

FMNH293240 (reproduced from Coles and Walsh, 2006, fig. 35°), distal genitalia and penial sections. 18. Same, opened to show

pilasters (reproduced from Coles and Walsh, 2006, fig. 36°), 19. D. dorfe uilliana, UF 376768 (second specimen), apical penis opened

to penial retractor muscle to show the terminal end of the pilasters, with obsolete ene into the vas deferens and the terminal

penial papilla. 20. Same, view of papilla from the side showing stalked form. 21. D. dorfeuilliana, Faulkner County, 10 October 2005,

NMW.Z.2005.011.03298, showing enlarged proximal vas deferens (vd) and vagina (vg) approximately twice as long as diameter, 22.

Same, penis opened to show pilasters. 23. D. dorfeuilliana, Carroll County, 10/15 March 2005, UF 376770, distal genitalia showing

enlarged vas deferens. 24. Same, detail of penis. Abbreviations: ag, albumen gland; at, atrium: ep, epiphallus; fo, free oviduct; hd,

hermaphrodite duct; pa, penial papilla: p, penis; pr, penial retractor muscle; pt, prostate gland; s, spermatheca; sd, spermatheca

duct; u, uterus; vd, vas deferens; vg, vagina. ° Reproduced by permission of the American Malacological Bulletin.

regarded previously by us as a specific character (Coles (Quercus spp.)-hickory (Carya spp.) forest. However,

and Walsh, 2006). However, this was not evident for the D. lithica is found under stones and fallen logs and on

three specimens from Faulkner County (Figure 21), al- bare ground in open, dry areas where trees and under-

though these conformed in other respects to the descrip- bch have been cut and where regeneration by young

tion given here. oak, hickory and juniper (Juniperus ashei Buchholtz,

1930) is occurring. Daedalochila dorfeuilliana lives in

humid conditions under stones and fallen logs on undis-

Habitats: Geographic eS of Daedalochila turbed forest floor but can also extend into areas of scrub

lithica and D. dorfeuilliana, by Arkansas counties, are where trees have been cut and on exposed limestone

similar (Figures 48, 49). Both species were found in oak outcrops.

Figures 25-32. Shells of Daedalochila lithica (25, 27, 29, 30) and Daedalochila dorfeuilliana (26, 28, 31, 32,) from Arkansas. 25.

Apical view, D. lithica, paratype, UMMZ 205895, 8.0 mm diameter, six miles east of Mountain View, Stone County. 26. D

dorfeuilliana, FMNH 308109, 8.2 mm diameter, Buffalo National River, Tyler Bend, Searcy County. 27. Umbilical view, D. lithica

same as above; 28. D. dorfeuilliana, same as above; 29. D. lithica paratype, FMNH 256372, 7.8 mm diameter, six miles east of

Mountain View, Stone County. 30. D. lithica, paratype, FMNH 256372, 7.5 mm diameter (same lot as 29 above). 31. D. dorfeuilliana,

FMNH 308114, $.2 mm diameter, Dogwood Trail at Beaver Dam Overlook, Carroll County. 32. D. dorfeuilliana, FMNH 308119,

S.0 mm diameter, Buffalo National River, Rush Historic Area, Marion County. Scale bars: figures 25-28 = 2.0 mm; figures 29-32

= 0.5 mm. Arrows indicate the slightly immersed palatal lamella of D. lithica (Figures 29, 30) and the more deeply immersed palatal

lamella of D dorfeuilliana Figures 31, 32

Page 136 THE NAUTILUS, Vol. 120, No. 4

5.0 5.35 5.7 6.0 7.5 9.0 2.5 3.75 5.0

0.8 1.1 14° 0.5 0.65 08 0.2 0.35 0.5

45 46 47

Figures 33-47. Box plot diagrams of shell measurements and ratios: Daedalochila dorfeuilliana (n=441) (above): Daedalochila

lithica (n=427) (below). The 25th-50th percentiles of data fall within the boxes, the 10th—90th percentiles fall within the error bars,

and data outside these ranges are plotted individually. The median value is shown within each box, or if not shown. falls at the lower

25% (Figures 33, 39, 41 [D. dorfeuilliana], 45) or upper 50% (Figure 41 [D. lithica]) limit of the boyes. No significant difference was

indicated between the means of cor responding pairs of characters (P>0.05), Student's t values are given in parentheses. 33. Number

of whorls (5.32). 34. Diameter of shell, mm (6.89). 35. Height of shell, mm (11.46). 36. Body whorl height, mm (10.93). 37. Outside

diameter of aperture, mm (8.90). 38. Inside diameter of aperture, mim (7.19), 39. Width of aperture, mm (4.31), 40. height of parietal

tooth, mm (15.62). 41. width of parietal tooth, mm (8.55). 42. diameter of umbilicus, mm (5.12). 43. height of shell diameter/

diameter of shell (6.65). 44. number of whorls/diameter of shell (9.89). 45. inside diameter of apertureAvidth of aperture (3.66), 46.

width of parietal tooth/height of parietal tooth (5.21). 47. diameter of umbilicus/diameter of shell (12.63),

—_— SS SSS

G. E. Walsh and B. F. Coles, 2006

Page 137

Figures 48-49. Distributions of D. lithica (Figure 48) and D. dorfeuilliana (Figure 49) in Arkansas. Dots: counties represented

in this study; blackened: counties not represented in this study.

DISCUSSION

Comparison of the shells of Daedalochila lithica and D.

dorfeuilliana show few consistent differences between

them: only the depth of the palatal lamellae within the

aperture provides a differential specific character (Table

1). Specimens selected on this basis show consistent dlif-

ferences in their genital anatomy (Table 1) that are as

marked as between other members of the D. plicata

group that have distinct shell morphology. For example,

Daedalochila peregrina (Rehder, 1932) differs from D.

lithica in its penis being of approximately uniform width

throughout its length, an obsolete epiphallus, and pilas-

ters anastomosed (Coles and Walsh, 2006). The penis

ot Daedalochila bisontes Coles and Walsh, 2006, is simi-

lar in shape to that of D. lithica, but differs in the pilas-

ters being more abruptly developed in the mid penis

(Coles and Walsh, 2006). Other species in the group,

Daedalochila plicata (Say, 1821), Daedalochila fastigiata

(Say, 1829), Daedalochila jacksoni (Bland, 1866) and

Daedalochila troostiana (Lea, 1839), are distinct in the

highly developed apical pilasters or associated features

(Coles and Walsh, 2006). Thus, these new data support

Hubricht’s view that D. lithica is a taxon distinct at spe-

cific level from D. dorfeuilliana and other Polygyridae of

the USA.

Position of Daedalochila lithica and D. dorfeuilliana

in the D. plicata group: In our earlier study of the D.

plicata group (Coles and Walsh, 2006), we abandoned

the concept of Millerelix (Pratt, 1981b) and the subgen-

era Millerelix sensu stricto and Prattelix (Emberton,

1995) because of the lack of, or variability of, the

diagnostic generic and subgeneric features. In part, this

decision was made because the anatomy of D. dorfeuil-

liana as we described (Coles and Walsh, 2006), and con-

firmed here, disagreed with that described by Pratt

(19S1b), particularly with respect to the diagnostic ge-

neric features of Millerelix: an elongate penis (width/

length<0.06), and the presence of a pendant conical pro-

jection in the apical penis (Pratt, 198la,b; Emberton,

1995; Coles and Walsh, 2006). The dissections presented

here show that these features are variable between indi-

viduals for both D. lithica and D. dorfeuilliana. For ex-

ample, both species show considerable variability of pe-

nis width/length, and both possess a minute papilla in the

terminal penis. The papilla was not found in all dissected

specimens, but because of the fragility of the tissues of

the epiphallus/terminal penis, we cannot be certain

Table 1. Consistent differences between Dacdalochila lithica and Daedalochila dorfeuilliana.

Feature D. lithica

D. dorfeuilliana

Palatal lamella of aperture not immersed (Figures 29, 30)

Penis approximately uniform in diameter from base

to mid length (Figures 5, 12, 14-16)

Proximal vas deferens

Vagina length/diameter

not greatly enlarged, approximately twice its

minimum diameter (Figures 6-13)

approximately 1 (Figures 12, 14-16)

iminersed (Figures 31, 32)

widest at mid length where the pilasters are

well developed, with a narrow base (Figures

17, 21, 24)

greatly enlarged, approximately 4 times its

minimum diameter (Figures 21, 23)

approximately 2 (Figure 21)

Page 138

THE NAUTILUS, Vol. 120, No. 4

whether it is variable in development or was destroyed

during dissection. Nevertheless, its presence agrees with

Pratt’s description of the anatomy of D. dorfe ruilliana

(Pratt, 1981b) and appears to be a “derivative” of the

apical, pendant, conical projection (Emberton, 1995). It

was not observed by us previously because several as-

pects of Pratt’s (198la, b) description discussed in Coles

and Walsh (2006) led us to believe that this feature was

more prominent than it is. It should also be noted that

Pratt (1981a, b) regarded the epiphallus (of D. lithica?)

as that portion of the narrow terminal penis extending

from the papilla to the penial retractor muscle (i-e., being

approximately 1/10 the length of the penis), whereas we

have regarded the epiphallus as the abrupt penial nar-

rowing (e.g g., Figure 24) observed in D. dorfeuilliana (i.e.,

the papilla is in the epiphallus). ). Other species of the D

plicata group were examined for papillae or similar fea-

tures through the opened terminal penis (i.e., to the pe-

nial retractor muscle) but none was observed (Coles and

Walsh, 2006). Using Emberton’s (1995) classification of

Millerelix and the anatomy as described by us, D. lithica

would be placed in the subgenus Millerelix on the basis

of the elongate, narrow penis and the lack of a greatly

enlarged proximal vas deferens, whereas D. dorfeuilliana

would be P laced in the subgenus Prattelix because of the

wider penis and greatly enlar ged vas deferens (Ember-

ton, 1995). Thus, these data give some support to the

concepts of Millerelix as ised! by Pratt (19Sla,b) and

modified by Emberton (1995), but they do not change

our conclusions concerning the inappropriate use of

Millerelix for all members of the D. plicata group.

ACKNOWLEDGMENTS

We are grateful to Jochen Gerber for providing shells

from the ‘Hubrieht collection deposited at FMNH, Liath

Appleton for providing shells a UMMZ, and to the

staff of the Buffalo National River, Greg Butts of the

Arkansas State Parks, Douglas Zollner of the Arkansas

Field Office of the Nature Conserv ancy, Cindy Osbome

of the Arkansas Natural Heritage Commission, the statf

of the Arkansas Game and Fish Commission and the staff

of the Ozark Saint Francis and Ouachita National For-

ests.

LITERATURE CITED

Branson, B. A. 1970. Shell oe in Polygyra dorfeuilliana.

The Nautilus $3: 120-13.

Coles, B. F. and G. E. Walsh. 2006. Daedalochila sp. nov. from

northwest Arkansas, U.S.A., the anatomy of the Polygyra

plicata group, and the validity of the genus Millerelix Pratt,

1981 (Gastrocopta: Pulmonata: Polygyridae). American

Malacological Bulletin 21: 99-112.

Emberton, K. C. 1995. When shells do not tell; 145 million

years of evolution in America’s polygyrid land snails, with

a revision and conservation properties. Malacologia 37:

69-110.

Hubricht, L. 1961. Eight new species of land snails from the

southern United States. The Nautilus 75: 26-33.

Hubricht, L. 1985. The distributions of the native land mollusks

of the eastern United States. Fieldiana, Publication Num-

ber 1359, The Field Museum of Natural History, Chicago.

viii + 191 pp.

Pilsbry, H. A. 1903. Mollusca of western Arkansas and et

states, with a revision of Paravitrea. Proceedings of the

Academy of Natural Sciences, Philadelphia. February

1903: 193-214.

Pilsbry, H. A. 1940. Land Mollusca of North America (North of

Mexico). epee: of Natural Sciences, Philadelphia,

Monogr aph L (2): 575-994 + ix pages.

Pilsbry, H. A. and J. a Ferriss. 1906. Mollusca of the Ozarkian

fama. Proceedings of the Academy of Natural Sciences,

Philadelphia. December 1906: 529-567.

Pratt, W. L. 1981a. A revision of the land snail genus Polygyra

in Texas. Unpublished Ph.D. dissertation, University of

Arizona, Tucson.

Pratt, W. L. 1981b. A revision of the land snail genus Polygyra

in re Dissertation Abstracts lnterstional 42: 1352-B.

Walsh, E. and B. F. Coles. 2002. Distributions and geo-

ete ae itionships of the polygyrid land snails (Mol-

lusca, Gastropoda, Polygyridae) of Arkansas. Proceedings

of the Arkansas Academy of Science 56; 212-219.

THE NAUTILUS 120(4):139-149, 2006

nga 7236

| age 39

The Neogene history of Pr isogaster Morch,

‘Tarbinidae) in South America

Thomas J. DeVries'

Burke Museum of Natural History

and Culture

University of Washington

Seattle, WA 98195 USA

1850 (Gastropoda:

ABSTRACT

Prisogaster Mérch, 1850, is the sole genus of the turbinid sub-

family Prisogasterinae Hickman and McLean, 1990, and is re p-

resented by only one extant species, the western South Ameri-

can P. niger (Wood, 1828). Two new fossil species from south-

ern Peru (P. valenciai new species and P. meleani new species)

extend the record of Prisogaster to the middle late Miocene

and reveal its turbinine ancestry. A major morphological trans-

formation during the late Pliocene produced the modern taxon,

which is heavier and more streamlined than its Mio-Pliocene

forebears, and thus probably better suited to the higher energy

environments that characterize the present-day Peruvian and

Chilean coastline.

INTRODUCTION

Prisogaster Moérch, 1850, has rightly been called “enig-

matic” by Hickman and McLean (1990) in their study af

trochoidean systematics. The genus has only one species

the extant Prisogaster niger (Wood, 1828), a gastropod

with a sturdy purple -black shell and calcareous opercu-

lum. The species is endemic to western South America

and has a fossil record not older than the middle Pleis-

tocene [Herm, 1969; not late Pliocene (Hickman and

McLean, 1990)]. Individuals of Prisogaster live amongst

wave-battered intertidal rocks and tide pools (Marin-

covich, 1973; Guzman et al., 1998), niches more typically

occupied by trochids than turbinids (Hickman and

McLean, 1990). Its perplexing suite of characters has

led Prisogaster to be placed in Phasianellinae Swain-

son, 1840 (Thiele, 1929: Wenz, 1938), Turbininae

Re ifinesque, 1S15 (Knight et al., 1960), and, most re-

cently, Prisogasterinae ( (Hiclanan and McLean, 1990).

This paper describes two new fossil species of Priso-

gzaster from southern Peru, P. stucchii new species and P.

valenciai new species. Specimens of the former were

found near Sacaco in middle upper Miocene and lower

Pliocene well-sorted cross-bedded sandstones. Speci-

' Mailing address: Box 13061, Burton, WA 98013 USA

mens of the latter were encountered farther south near

Chala in upper lower Pliocene or upper Pliocene bio-

clastic gravels. These new taxa demonstrate a dramatic

shift in morphology and niche of the living species, P.

niger, from its Mio-Pliocene prede cessor.

GEOLOGY

The Neogene stratigraphy of forearc basin deposits be-

tween Pisco and Camana (Figure 1) was reviewed by

DeVries (1998). Upper Miocene and Pliocene marine

strata of the Pisco and La Planchada formations include

fine-grained tuffaceous and diatomaceous sandstone,

which are attributed to outer shelf environments, and

coarse-grained massive tuffaceous sandstone from closer

to shore (Muizon and DeVries, 1985). Lying disconform-

ably on crystalline basement rocks are cross-bedded and

lenticular bioclastic conglomerates, remnants of littoral

Paracas

Peninsula &

[2 Pisco Basin

0 100 km

Figure 1. Location of the Pisco forearc basin in southern

Peru. New fossil species of Prisogaster are from Cenozoic de-

posits near Sacaco and Chala.

Page 140

THE NAUTILUS, Vol. 120, No. 4

deposits that lapped onto pre-Eocene erosional plat-

forms or against precipitous Andean foothills. An excel-

lent example of the latter is seen southeast of Chala,

where the Panamerican Highway descends in tight

curves towards the beach at Playa Huacllaco (Figure 2).

Seventy meters of sediment (Figure 3) were de ‘posite din

high-energy foreshore and intertidal environments that

once flanked rugged cliffs (DeVries, 2003). The age of

the Huacllaco be ds is constrained by b basal beds “with

specimens of Concholepas nodosa Hupé, 1854, Acan-

thina triangularis DeVries, 2003, and Herminespina

mirabilis (Moricke, 1896), which collectively indicate an

early late Pliocene age (DeVries and Frassinetti, 2003),

and the uppermost and oldest of several marine terraces,

whose 200 m elevation and largely extant taxa suggest a

latest Pliocene age (Muizon and DeVries, 1955).

MATERIALS AND METHODS

Specimens described in this study were found by the

author. C sompar ative material was studie dat the Los An-

ALTO GRANDE©

DV 571-1

Aguada

de Lomas

15°30'S

Contour interval is 100 m

PANAMERICAN

HIGHWAY

@ Locality

samples

geles (California) County Museum of Natural History

LACM). Locality and sample descriptions are listed in

the appendix. * DV” locality numbers refer to the author's

field notes. Lengths (L) and widths (W) are measured in

millimeters. Dimensions of broken specimens are en-

closed by parentheses. Figured specimens viewed from

wey angles may be listed with a “maximum viewed

width” (m.v. width) measured at right angles to the axis.

Some figured specimens are coated with ammonium

chloride. Types and numbered specimens are deposited

at the Departamento de Paleontologia de Vertebrados,

Museo de Historia Natural, Universidad de San Marcos,

in Lima, Peru (MUSM INV) and University of Washing-

ton’s Burke Museum of Natural History and Culture in

Seattle, Washington (UWBM).

SYSTEMATICS

Family Turbinidae R Rafinesque, 1S15

Subf amily Prisogasterinae Hickman and McLean, 1990

—

}

“DV 1029-1,

DV 460-1,

15°40'S

a 1635-1

SCALE

Contour interval is 50 1m

2km

SCALE

Contour interval is 100 m Playa Huacllaco

** Includes DV 1254-Bal 6, -Bal 8, and -Bal 10 (see Appendix)

Figure 2. Type localities of Prisogaster valenciai new species (locality-samples DV 1254-Bal 6, Bal 8, and Bal 10) and P. stucchii

new species DV 571-1

and Sacaco (DV 380-2, DV 573-1) with

P. niger (DV 1629-1]

Also shown are lower Pliocene locality-samples near Yauca (DV 1029-1, DV 1635-1, DV 460-1, DV 1598-1)

P. meleani and an uppet Pliocene/lower Pleistocene marine terrace southeast of Chala with

T. ]. DeVries, 2006

Section above

Playa Huacllaco

l—=DV 1629-1

unit] /0 Te ae

Face (P. niger)

— 3

2 — DV 1254-Bal 10

(P. valenciai)

; DV 1254-Bal 8

40 — los were (P. valenciai)

ot DV 1254-Bal 6

= (P. valenciai)

—

—| 30-4

20 -

Zz [

10 -

meters eeseasies

& | Cobbles Massive sandstone

j ; Foncarnl ps Pee

} | Pebbly sandstone +) Bioclastic rippled sandstone

f Wy a p .

| Polychaete colony -») Bioclastic crossbedded sandstone

Es Sn - Fee

B. ‘| Coquina J3.234 Igneous basement

Figure 3. Huacllaco section southeast of Chala with strati-

graphic position of type specimens of P. valenciai new species

and other horizons with Prisogaster material.

Genus Prisogaster Mérch, 1850

Amyxa F. H. Troschel, 1852 (objective synonym)

Type species: Turbo niger Wood, 1828. Type locality

not specified.

Discussion: Moérch (1850, p. 21) did not describe

Prisogaster, but simply applied the new genus name

without comment to Turbo niger Gray (= = Turbo niger

Wood, 1828), itself todeed with only a drawing aoa

one-word description, “black” (Wood, 1828, p. 18). Hick-

man and McLean (1990) imp licitly defined Prisogaster

by their description of Prisogasterinae. Diagnostic non-

anatomical characters faclade an incomplete peristome

(also present in Turbininae) and an operculum with a

“thick, convex exterior calcareous pad” (Hickman and

McLean, 1990, p. 52). Other distinguishing shell charac-

ters included the black color of the outer shell layer, an

oblique aperture, the absence of an umbilicus in adult

specimens, a predominance of spiral sculpture, and an

inner nacreous layer. New fossil data show that the black

color is not diagnostic of the entire genus. The morphol-

ogy of the operculum is distinctive for the subfamily and

genus, however, as is the quadripartite structure of the

columella described in this paper.

Prisogaster niger (Wood, 1528)

(Figures 4-10, 13, 14)

Turbo niger Wood, 1828, p. 18, pl. 6, fig. 1; @Orbigny, 1840, 5:

411-412, vol. 9 (Mollusca), pl. 55, figs. ae

Turbo niger Gray, 1839, p. 143, pl. 36, fig. 1; Hupeé, 1554, p.

140.

Turbo (Prisogaster) niger Wwood.—Dall, 1909, p. 235.

Prisogaster niger Wood.~-Morch, 1850, p. 21; Carcelles and

Williamson, 1951, p. 268; Dell, 1971, p. 197; Marincovich,

1973, p. 24, fig. 41; Osorio et al., 1979, p. 1S, fig. 15;

Ramirez, 1981, p. 130, fig. 154; Alamo and Valdivieso,

1997, p. 15, fig. 30; Guzman et al., 1995, p. 37, fig. 27;

Forcelli, 2000, p. 64, fig. 102; Aldea and Valdovinos, 2005,

p. 390, fig. S-H.

Prisogaster niger minor Mérch, 1850, p. 21.

Trochus gaudichaudii Hupé, 1854, p. 146, Malacologia, pl. 4,

figs. 4, 4a, 4b.

Diagnosis: Spiral cords and interspaces purple-black.

Whorls slightly angulate anteriorly. Spiral sculpture of 13

to 24 primary spiral cords, including six to ten cords on

base.

Description: Shell thick, up to 30 mm long, globose,

variably compressed avially. Spire variably elevated:

about one-quarter to one -third of shell length. Proto-

conch unknown; teleoconch with about five whorls.

Whorls broadly rounded posteriorly, very weakly angu-

late or biangulate anteriorly; periphery anterior to aia!

midpoint of body whorl. Sutures usually appressed,

sometimes slightly impressed. Exterior purple black,

spire usually corroded, nacreous. Axial sculpture absent.

Spiral sculpture of 13 to 24 primary spiral cords, includ-

ing six to ten on base; cord widths vary irregularly two-

fold. Most posterior spiral cord wider, flatter, forming

collar against preceding whorl. Interspaces narrower

Page 142

THE NAUTILUS, Vol. 120, No. 4

white band 1 O

14 15

Figures 4-15. Prisogaster spp. 4-10. Prisogaster niger (Wood, 1828). 4. UWBM 97829, Pisco Bay, Recent, abapertural view,

length = 18.4 mm. 5. UWBM 97830, Pisco Bay, Recent, oblique basal-apertural view, m. v. width = 15.7 mm. 6. MUSM INV 116,

DV 1629-1, latest Pliocene, oblique basal- apertural view showing quadripartite structure of columella, m. v. width = 17.9 mm. a =

inner oe column, b = medial nacreous band, c = outer white sinuous ridge, d = outermost elongate excavation, i = inner nacreous

layer, ii = middle shell layer, iii = outer purple-black calcitic layer

7. UWBM 97829, basal view, width = 19.1 mm. 8. UWBM 97837

DV 1699- 1, operculum, interior view, length = $.9 mm. 9. UWBM 97837, Ss exterior view. 10. UWBM 97828, DV 398-1,

Recent, apertural view showing outer layer and inner nacreous layer on body whorl, length = 18.3 mm. 11, 12. Prisogaster mcleani

new species. Early Pliocene. 11. UWBM 97548, DV:

3-1. operculum, interior view, length = = 8.7 mm. 12. UWBM 97848, exterior

view. 13, 14. Prisogaster niger (Wood, 1828), 13. UWBM 97831, DV 461-1, latest Pliocens. basal view, m.v. width = 18.9 mm. 14.

UWBM 97831, abape rtural view, length =

DV 1254-Bal 8, apertural view, width = 15.9 mm.

than spiral cords, shallow, sometimes crossed by strongly

prosocline colabral growth-line lamellae. Aperture ob-

lique, diagonally ovate, with incomplete peristome. Um-

bilicus absent in juveniles and Sls Outer lip thick,

strongly prosocline, with weak inflection posteriorly and

16.3 mm. 15. Prisogaster valenciai new species. Late Pliocene. UWBM 97838, holotype,

salient adjacent to suture; inner edge smooth, nacreous.

Parietal and umbilical areas weakly excavated, nacreous;

sometimes with weak anal canal without sinus. Columella

with innermost white column ending anteriorly at slight

inflection near columellar base; me sdially with slightly ex-

T. J. DeVries, 2006

cavated, curved, nacreous band alongside innermost col-

umn, narrowing anteriorly, then broadening at base of

aperture to join nacreous inner layer of outer lip; also

with weakly sinuous white ridge outboard of nacreous

band, extending to base of aperture and coalescing with

interspace between second and third basal spiral cord to

form a blunt tooth; first tooth separated from second

blunt tooth abaxially by short groove inside aperture; and

with shallow, white, elongate excavation bordering ridge

at outer margin of columellar area.

OPERCULUM: Exterior convex, steeper posteriorly and

adaxially; basal rim smooth, remainder of surface pustu-

lose, more so peripherally than centrally. Two subaxially

elongate, adaxially- anteriorly converging creases on

adaxial side of diz wonal bulge, inner crease extending to

abaxial posterior corner; sinuous crowth lines present on

abaxial side. Interior nearly planar, coiled, changing from

multispiral to paucispir al: all coils with long growing

edge. Subsurface texture reticulate with elongate ‘ “cells”

pe rpendicular to growth lines.

Material Examined: MUSM INV 113, DV 461-1, lat-

est Pliocene, L 13.6, W 15.6; MUSM INV 114, DV 464—

1, middle Pleistocene, L 22.8, W 22.0: MUSM INV 115,

DV 463-1, late Pleistocene, L (21.7), W 25.7; MUSM

INV 116, DV 1629-1, latest Pliocene, L 15.3, W 17.9:

MUSM INV 117, DV 1629-1, L 12.7, W 14.1; UWBM

97828, DV 398-1, Recent, L 18.3, W 17.4; UWBM

97829, Pisco Bay, Recent, L 15.4, W 19.1; UWBM

97830, Pisco Bay, Recent, L 14.6, W 15.1; UWBM

97831, DV 461-1, L 16.3, W 18.5: UWBM 97832, DV

464-1, L 21.0, W 21.6; UWBM 97833, DV 463-1, L

(23.5), W 27.0: UWBM 97834, DV 720-1, Holocene,

operculum, L5.1, W 4.4; UWBM 97835, DV 1629-1, L

16.0, W 19.1; UWBM 97836, DV 1629-1, L 13.5, W 14.2;

UWBM 97837, DV 1629-1, operculum, L 8.9, W 7.1.

DeVries collection: Ipun, Chile, Recent, lot of one;

Caldera region, northern Chile, from eight meters deep

on sand and rocks, Recent, lot of two: DV 461-1, lot of

two: DV 463-1, lot of six: DV 720-1, lot of twelve; DV

730-1. Holocene, lot of one and one operculum; DV

1252-1, latest Pliocene, lot of 3.

Distribution: Ecuador (7°N) to northern Peru:

LACM collections, Recent. North-central Peru: Ho-

locene, Recent. Southern Peru: latest Pliocene, early

Pleistocene, middle Pleistocene, late Pleistocene, Re-

cent. Chile: middle Pleistocene to Recent (Herm, 1969;

Valdovinos, 1999).

Remarks: Modern specimens of Prisogaster niger vary

in the number of spiral cords and height of the spire. The

oldest specimens of P. niger from southern Peru have

only 13 to 15 primary spiral cords between the columella

and suture (e. s.. Figures 13, 14), about as few as the

youngest specimen of the older P. valenciai. Specimens

of P. niger differ from those of P. valenciai by lacking any

vestige of cream-colored banding of cords or interspaces

The moder range of Prisogaster niger extends from

7° N to 41° S (LACM collections) and beyond to the

Straits of Magellan (Osorio et al., 1979; Valdovinos,

1999), a Gistuhion more extensive than ascribed to the

species by Dall (1909) or Alamo and Valdivieso (1997).

The Pleistocene record is limited to southern Peru and

Chile, with no records from the marine tablazos of north-

em Peru (DeVries, 1986). The late Pliocene record is

still more restrictive, with only southern Peruvian speci-

mens known, including the oldest, which were found

southeast of Chala (DV 1629-1) in the uppermost co-

quina (Unit TV) of the Huacllaco section (Figure 3), a

shell bed dominated by thick disarticulated valves of the

bivalve, Mulinia edulis (King, 1831).

A second modern species assigned to Prisogaster has

been the Chilean P. elevatus (Eydoux and Souleyet,

1852) (Souleyet, 1852, v. 2, p. 594, pl. 37, figs. 15-19;

Nicosia and Gaete, 2003). The original figures of “Turbo

elevatus,’ however, show a specimen with a blotchy

purple-black color, elevated spire, absence of spiral

sculpture, wavy growth lines, and evenly convex calcar-

eous operculum visibly coiled on both sides, all features

characteristic of Tricolia Risso, 1826 (Hickman and

McLean, 1990), more particularly the Chilean Tricolia

meleani Marincovich, 1973. Neither the 14 mm le neth of

Eydoux and Souleyet’s “Turbo elevatus,” howeven nor

the deep lunate columellar/parietal excavation, is char-

acteristic of Chilean Tricolia, but rather suggest the Chil-

ean trochid, Diloma nigerrima (Gmelin, 1791). The ge-

neric assignment of “Turbo elevatus” remains in doubt.

Prisogaster valenciai new species

(Figures 15-19, 22)

Diagnosis: Compressed axially: sculpture of five to 14

broadly rounded charcoal-gray primary spiral cords, in-

cluding two to four on the b vase; One or more interspaces

cream- “60108 ed.

Description: Shell less than 15 mm long, globose,

compressed axially. Thickness indeterminate (inner shell

layers missing). ae moderately elevated, length inde-

terminate (much of spire missing). Protoconch and early

whorls of teleoconch unknown. Whorls broadly rounded

posteriorly, weakly angulate anteriorly. Periphery usually

anterior to axial midpoint of whorl. Sutures slightly im-

pressed. Exterior gray to black, with interspaces some-

times cream- “colored. Axial sculpture absent. Spiral

sculpture of five to 14 broadly rounded primary cords,

three to LO posterior to base, two to four on base; spiral

cords about equally wide except for broader, flatter, most

posterior spiral cord forming low collar adjacent to su-

ture. Interspaces usually narrower than spiral cords,

sometimes filled with single secondary spiral cord.

Strongly prosocline growth lines, sometimes lamellate.

Aperture oblique, diagonally ovate. Outer lip strongly

prosocline. Columella and umbilical area missing. Oper-

culum unknown.

Type Material: UWBM 97835, DV 1254-Bal 8, holo-

type, late Pliocene, L (13.3), W 15.9: UWBM 97539,

paratype, DV 1254-Bal 6, L (13.8), W 15.5; MUSM INV

Page 144 THE NAUTILUS, Vol. 120, No. 4

T. J. DeVries, 2006

Page 145

L1S, paratype, DV 1254-Bal 8, fragment; MUSM INV

119, paratype, DV 1254-Bal 10, fragment.

Type Locality: Roadcut along the Panamerican High-

way, 10 km southeast of Chala. Locality-samples DV

1254-Bal 6, DV 1254 Bal-S, DV 1254-Bal 10, late

Pliocene, in section at 35, 42, and 47.5 meters, respec-

- ely, above crystalline basement (Figures 2, 3), 15°52’

_ 74° 10' W (Chala 1:100,000 qui adrangle).

Other Material Examined: UWBM 97840, DY

LO31-1, late early Pliocene, L (12.2), W 13.0.

Distribution: Southern Peru: late early Pliocene to

late Pliocene.

Etymology: Named in honor of Dr. Niels Valen-

cia C ee director of the Museo de Historia Natural,

Universidad Nacional Mayor de San Marcos, Lima,

Peru.

Remarks: The incomplete specimens of Prisogaster

valenciai preclude a complete description, but the shape,

size, and spiral sculpture clearly mark them as exam-

ples of Prisogaster. These specimens differ from P. niger

a having one or more cream-colored interspaces and

gene rally fewer and more clearly differentiated pri-

mary spiral cords. The oldest known specimen of P.

valenciai is from the Pliocene section above Playa

Huacllaco (Unit I, DV 1254-Bal 6): it has several broad

dark spiral cords alternating with cream-colored bands

that sometimes aoe a spiral cord and interspace

(Figures 1S, 22). A specimen from several kilometers

away (DV 1031-1) has very few spiral cords, all gray, and

equally wide interspaces, all cream-colored (Figures 16,

17, 19).

Most specimens of Prisogaster valenciai were found in

Unit II and the lower part of Unit III of the Huacllaco

section (DV 1254; Figure 3) in cobbly barnacle-rich bio-

clastic gravel and poorly sorted coarse-grained sandstone

thought to have been deposited intertidally or at very

shallow subtidal depths. Associated molluscan taxa in-

clude Acanthina spp: Concholepas spp.; Choromytilus

chorus (Molina, 1782); and a new species of Xanthocho-

rus Fischer, 1884 (DeVries, 2005a).

Prisogaster mceleani new species

(Figures 11, 12, 20, 21, 23-32)

Diagnosis: Globose, whorls broadly rounded anteri-

orly and posteriorly; tan-colored, often mottled, striped,

or speckled with brown.

Description: Shell globose, about 20 mm long. Spire

moderately to greatly elevated, about 30% to 40% of

shell length. Protoconch unknown, teleoconch with

about ane whorls. Whorls convex, broadly rounded pos-

teriorly and anteriorly, without anterior angul: ations. Su-

tures variably impre ssed. Exterior tan, often with mot-

tling, prosocline flammules, or zig-zag patterns of brown.

Axial sculpture absent. Spiral sculpture of ten to 15

rounded primary spiral cords between suture and um-

bilical area, subdued on smooth specimens; secondary

spiral cords rarely interspersed. Entire surface often w ith

tertiary spiral threads. Interspaces variably wide, crossed

by strongly prosocline, colabral growth-line lamellae. Ju-

veniles wath three weaker spiral cords on base, four to

five primary spiral cords on whorl. Aperture oblique,

ovate to nearly circular, peristome incomplete. Umbili-

cus absent. Outer lip thin, strongly prosocline, inner edge

smooth. Parietal and umbilical areas weakly excavated in

adults. Columella with inner white cohamn curving an-

teriorly; with a narrow, slightly excavated, nacreous hand

merging anteriorly with nacre-lined aperture; with a

white ridge, slightly sinuous, outboard of the nacreous

band, extending to base of aperture and coalescing with

interspace neta een second and third basal spiral coud,

with one or two short teeth abaxial; and with an outer-

most, narrow, white, excavation bordering the ridge at

the outer margin of the columellar area, extending ante-

riorly just over half the length of columella.

OPERCULUM: Exterior convex, steeper posteriorly and

adaxially; basal rim smooth, remainder of surface pustu-

lose, more so peripherally than centrally. Two sub-axially

elongate, anteriorly converging creases on adaxial side of

diagonal bulge, imner crease extending to abaxial poste-

rior corner; sinuous growth lines present on abaxial side.

Interior nearly planar, coiled, multispiral changing to

paucispiral; all coils with long growing edge. Larger coils

with centered shallow, iscoad. Sathottomed channel.

Subsurface texture reticulate with elongate “cells” per-

pendicular to growth lines.

Figures 16-32.

Prisogaster spp. 16-19. Prisogaster valenciai new species. Late Pliocene. 16. UWBM 97840, DV 1031-1, early late

Pliocene, oblique see view, width = 13.0 mm. 17. UWBM 97840, apertural view. 18. UWBM 97839, paratype, DV 1254-Bal 6,

abapertural fragment. 1

v. width = 12.7 mm. 19. UWBM 97840, oblique basal-apertural view, m. v. width = 12.4 min. 20, 21.

Prisogaster mcleani new ek 20. MUSM INV 121, DV 1598-1, early Pliocene, oblique spire view showing change in color pattern

and repaired break, m. v. width = 12.1 mm. 21. MUSM INV 121, ape rtural view showing naticid drillhole to left of columella, width

= 11.5 mm. 22. Prisogaster valenciai new species. Late Pliocene. UWBM 97839, oblique basal view, m. v. width = 12.7 mm. 23-32.

Prisogaster mcleani new species. 23. UWBM 97841, DV 571-1, holotype, middle late Miocene, apertural view showing quadripartite

structure of columella (see text), length = 13.9 mm. 24. UWBM 97842, DV 571-1, paratype, abapertural view, length = 21.1 mm

25. MUSM INV 120, DV 1595-1, apertural view showing quadripartite columella and prosocline color stripes, length = 12.4 mm. 26.

UWBM 97541. oblique spire view, m. v. width = 16.5 mm. 27. UWBM 97842, abapertural view. 28. UWBM 97844, DV 1598-1,

abape srtural view of juvenile showing bicarinate whorls and colabral growth-line lamellae, length = 9.4 mm. 29. MUSM INV 124, DV

573-1, early Pliocene, abapertural view showing flecked stripes, lensth = = 12.7 mm. 30. MUSM INV 125, DV 571-1, abapertural view

showing brown zig-zag pattern, length = 17.5 mm. 31. UWBM 97843, oblique spire view, m. v. width = 19.7 mm, 32. UWBM 97843

apertural view, length = 17.9 mm.

Page 146

THE NAUTILUS, Vol. 120, No. 4

Type Material: UWBM 97541, DV 571-1, holotype,

middle late Miocene, L 13.9, W 15.4: UWBM 97842, DV

571-1, paratype, middle late Miocene, L 21.1, W (18.5);

UWBM 97843, DV 571-1, paratype, L 17.9, W 19.5.

Type Locality:

intersection with abandoned paved road to San Juan de

Marcona (= El Jahuay locality of Muizon and DeVries,

1985). Shell banks on south- facing slope, 15°26'57" S,

74°52'06" W (Acarf 1: 100,000 quadr angle).

Material Examined: MUSM INV 120, DV 1598-1, L

12.4, W 12.1; MUSM INV 121, DV 1598-1, L 11.1, W

11.5; MUSM INV 122, DV 1635-1, early Pliocene, L 9.4,

W 11.0; MUSM INV 123, DV 460-1, lot of two; MUSM

INV 124, DV 573-1, L 12.7, W 14.5; MUSM INV 125,

DV 571-1, L 17.8, W 16.0: UWBM 97844, DV 1598-1,

early Pliocene, L 9.4, W 8.2; UWBM 97845, DV 1598-1,

L 12.1, W 12.7; UWBM 97846, DV 460-1, early

Pliocene, L 8.9, W 10.3; UWBM 97847, DV 573-1, early

Pliocene, L 12.0, W 13.0; UWBM 97848, DV 573-1,

operculum, L 8.7, wi S$. DeVries collection: DV 1598-1,

lot of four; DV 423-3, middle Pliocene, lot of two; DV

1029-1, early Pliocene, lot of two; DV 806-1, late Mio-

cene, lot of one; DV 380-2, early Pliocene, lot of one.

Distribution: Southern Peru: middle late Miocene,

early Pliocene.

Etymology: Named after James H. McLean, mala-

cologist, Natural History Museum of Los Angeles County.

Remarks: Specimens of Prisogaster stucchii differ

from those of P. niger by being uncompressed axially, by

having an outer caleitic layer that i is tan, sometimes with

brown mottling and prosocline axial stripes, and by hav-

ing spiral cords that are narrower and higher, with

broader interspaces. The complex structure of the col-

umella, however, is identical with that of P. niger, as is

the operculum. Prisogaster mcleani exhibits considerable

variation in spiral ornamentation, with specimens from

the same horizons being alternatively densely sculptured

with 15 spiral cords (Figures 24, 30), moderately sculp-

tured with eight primary and two secondary cords (Fig-

ure 26), and Tightly sculptured with ten subdued cand

that barely Hse Aue e the neighboring wide inter spaces

(Figures 31, 32). Some Miocene and lower Pliocene ju-

ve ail eames ns (Figures 28) have a pronounced bian-

gulate profile produced by two strong primary spiral

sores posterior to the suture, a feature shared with ju-

veniles of many turbinine species (Hickman and

McLean, 1990).

The oldest specimens of Prisogaster mcleani were

found near Alto Grande (DV 571-1) in middle upper

Miocene beds with an estimated age of about 9 Ma

(Muizon and DeVries, 1985). Lower Pliocene specimens

from Sacaco (DV 380-2) and Yauca (DV 1598-1, DV

1635-1) are identical with the Miocene specimens.

DISCUSSION

The extant Prisogaster niger, which appeared about two

million years ago, has changed little in either its pattern

Alto Grande, about one km south of

of low rounded spiral cords or its uniformly purple-black

color. Prisogaster mceleani, in contrast, resident on south-

ern Peruvian coasts from 9 Ma to 3 Ma, has highly vari-

able spiral sculpture and coloration. Some specimens

(Figures 24, 28, 30) exhibit the high spires, impressed

sutures, numerous primary spiral cords, and bicarinate

juvenile whorls that characterize many members of Tur-

bininae. These similarities strengthen the case of Hick-

man and McLean (1990) that Prisogasterinae and Tur-

bininae are sister taxa and that Prisogaster arose from an

advanced turbinine.

Prisogaster probably appeared in southern Peru dur-

ing the early late Miocene, since no turbinines have been

founeli in middle Miocene or older beds in southern Peru.

Its arrival coincided with that of other taxa that consti-

tuted the core of a late Miocene-early Pliocene mollus-

can fauna in southern Peru and northern Chile (DeVries,

2002). The route by which Prisogaster or its predecessor

arrived in Peru is unclear. No medium-sized or large

turbinids are known from Miocene deposits of Chile

(Philippi, 1587; Tavera, 1979; Nielsen et al., 2004). The

only Miocene turbinid from northern Peru (Spieker,

1922) is probably related to two northern Peruvian Re-

cent species of Turbo (Taeniaturbo) Gray, 1850; the

modern Patty have opercula with strong and complex

external spiral relief very unlike the sculpture on the

opercula of fossil and Recent species of Prisogaster. The

pustulose convex opercula and multipartite dalumellae of

some species of Turbo (Marmarostoma) Swainson, 1829,

do resemble those of Prisogaster, although opercula of

the former are more broadly convex and not scored by

diagonal grooves, and the columellae of the shell do not

have a well differentiated ridge and groove structure to-

ward the perimeter of the columellar area. Modern spe-

cies of Turbo (Marmarostoma) have an Indo-Pacific dis-

tribution, suggesting that Prisogaster might be a trans-

Pacific immigrant, as other western American taxa have

been (Emerson, 1978), including, it appears, the trochid

species, Diloma nigerrima (Gmelin, 1791) (Donald et al.,

2005).

The transition from the globose tan-colored and

brown-mottled Prisogaster mcleani to the axially com-

pressed purple-black P. niger occurred during the time

represented by Units I] and III of the Huadllaco section

(Figure 3), i.e., latest early Pliocene to late Pliocene, in

the guise of P. valenciai. The oldest specimen of P. va-

lenciai (Figures 18, 22; DV 1254-Bal 6) show the first

appearance of broad black spiral cords and the persis-

tence of thin cream-colored interspaces, the latter of

which become relegated to spire whorls on the youngest

specimens (Figure 15; DV 1254-Bal 10).

The late Pliocene appearance of a purple-black exter-

nal calcitic layer in Prisogaster is an odd event that none-

theless is repeated in western South American Diloma

Philippi, 1845 (Marincovich, 1973; Nielsen, 2003). The

significance of these purple-black outer layers in Quater-

nary turbinids and western South American trochids

[Miocene-Pliocene undescribed species of Tegula (Chlo-

T. ]. DeVries, 2006

rostoma) Swainson, 1840; Tegula (Chlorostoma) atra

(Lesson, 1830); T. ) luctuosa (VOrbigny, 1541); T.

(C.) tridentata (Potiez and Michaud, 1838); and, incipi-

ently, in T. (C.) quadricostata (Gray, 1828)], as well as

trochoids from mid- and high- latitude northern Pacific

and South African shorelines, merits further inve stiga-

tion.

Occurrences of most modern and Pleistocene speci-

mens of Prisogaster niger are consistent with a high-

energy inte srtidal and shallow subtidal habitat with Hele

and-gravel substrates (e.g., DeVries collection, Recent,

juv eniles, eight meters deep, northern Chile; DV 720-1,

Holocene, cravel beach ridge, northern Peru; DV 1629-

1, upper Pliocene, 250 m marine terrace, southern Peru).

The same is true for upper Pliocene occurrences of

specimens of P. valenciai, which are found in bioclastic

coarse-grained sandstones and gravel deposited within

150 meters of a mountainous paleo-shoreline. In con-

trast, specimens of the older P. mcleani are found in

well-sorted hummocky cross-bedded sandstones with

shell banks of large venerid bivalves, as well as bioclastic

sandy gravels, suggesting that the habitat of the late Mio-

cene- early Plosene species also included foreshore en-

vironments with lower energies than typical for intertidal

environments.

The late Pliocene was a time of morphological novelty

for Mio-Pliocene taxa in Peru other than Prisogaster:

Acanthina Fischer von Waldheim, 1807 (DeVries, 2003),

Concholepas Lamarck, 1799 (DeVries, 2000), Xantho-

chorus Fischer, 1884 (DeVries, 2005a), and Tegula Les-

son, 1835 (DeVries, unpublished data). It was during this

time that the molluscan fauna of the Mio-Pliocene Pe-

ruvian Faunal Province was undergoing the second

phase of a local species-level mass extinction (DeVries,

2001) that coincided or at least overlapped with in-

creased rates of tectonic uplift in northern and southerm

Peru (DeVries, 1986, 1988: Ortlieb et al., 1995) and the

transport of greater quantities of gravel and rounded

cobbles of f Andean andesitic rock to the southern Peru-

vian coast (DeVries, 2003).

CONCLUSION

With the discovery of two new fossil species in southern

Peru. the “enigmatic” turbinid Prisogaster now has a

pedigree extending back to the middle late Miocene.

Miocene and early Pliocene specimens with turbinine

features suggest Prisogaster did arise from a middle Mio-

cene species of Turbo, a possibility raised by Hickman

and McLean (1990). The place from which Prisogaster or

its turbinine ancestor immigrated is uncertain, hae based

on some similarity in key characters with some species of

Turbo (Marmarostoma), the Indo-Pacific region seems

as likely as the Magellanic or Panamic regions. The Priso-

gaster shell underwent a rapid transformation during the

late Pliocene, including an axial compression, a stream-

lining of sutural contacts, a thickening of the shell,

broadening and smoothing of primary spiral cords, and

the development of a purple-black outer shell layer.

Some of these changes might be construed to have im-

proved the strength of Prisogaster shells to more suc-

cessfully Gnihstand attacks from clawed predators, which

seem to have been a common hazard for individuals of P.

mceleani (see Figures 20, 24, 30, 31). Alternatively, a

stronger and more stable shell may have enabled indi-

viditals of P. niger to survive on rockier, higher-energy,

intertidal sibstates Such environments letans the

norm for coastal Peru when much of the coast com-

menced a 200 m uplift during the latest Pliocene and

protected embayments became fewer and much smaller

(DeVries, 2001).

ACKNOWLEDGMENTS

I would like to thank M. Urbina, R. Salas, and M. Stucchi

of the Departamento de Paleontologia de Vertebrados,

Museo de Historia Natural, Universidad Nacional Mayor

de San Marcos, in Lima, Peru, for their hospitality and

help in the laboratory and field. Helpful suggestions for

improving the manuscript were offered by C. Hickman

and S. Wielsen: Support for field research was provided

in part by a Fulbright scholarship in 1999.

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APPENDIX

Locality-samples:

DV 380-2 Sacaco, in cross-bedded eater just

low farmhouse, 15°32'29" S, 74°43'53"” W (Acari

100,000 quadrangle), Lower Pliocene.

DY 395-1 Playa Canastones, Bahia de la Independen-

cia, Peru (Punta Grande 1:100,000 quadrangle), Recent.

DV 423-1 Terrace de sposit capping eas strata

west of Quebrada Huaricangana, 14°55'29" 8S, 75°17'54"

W (Palpa 1:100,000 quadrangle), Pliocene.

DV 460-1 Roadcut along Panamerican Highway, de-

T. J. DeVries, 2006

P age 2 149

scent from north into Yauca. Shell beds, 15°39'49”" S

T4°31'50" W (Yauca 1:100,000 quadrangle),

Pliocene (same as DV 1598-1).

DV 461-1 Highest marine terrace about five km north

of Chala, Peru, uppermost Pliocene/lowermost Pleisto-

Lower a1

cene,

DV 463-1 Lower terrace, 5 km north of Chala (Chala

1:100,000 quadrangle), upper Pleistocene.

DV 464-1 Mid-level marine terrace, five kim north of

Chala (Chala 1: 100,000 quadrangle), middle Pleistocene.

DV 571-1 Alto Grande, about one km south of inter-

section with abandoned paved road to San Juan de

Marcona, on south-facing hillside west of Panamerican

Highway; one of several shell banks, 15°26'57” S,

74°52'06" W (Acari 1:100,000 quadrangle), middle upper

Miocene.

DV 573-1 Sacaco, shell banks and cross-bedded sand-

stone just below level of farmhouse (Acari 1:100,000

quacmenele), lower Pliocene.

DV 720-1 Eastern major cobble ridge near Santa pa-

leolagoun, northern Peru (see DeVries and Wells,

1990), Holocene.

DV 730-1 Between Ceros Pimenco and Mentiroso,

mouth of Santa paleo-lagoon, northern Peru (see

DeVries and Wells, 1990), Holocene.

DV 806-1 Southwest of Quebrada Usuaca, about 1.7

kan west of Hacienda Tunga building, in middle of small

natural amphitheater in hillside, highest shell bed. (Palpa

1:100,000 quadrangle), lower Pliocene.

DV 1029-1 Yauca Depression, west of Panamerican

Highway, 15°39'29" S, 75°35'08" W (GPS, Yauca 1:

100,000 quadrangle). lower Pliocene.

DV 1031-1 Section along Panamerican Highway,

Quebrada Huambo, about ten km west-northwest of

Chala. Roadcut along Panamerican Highway, north

face. 15°4S'41" S, 74°21'1S" W (GPS; Chala 1:100,000

quadrangle). Uppermost Pliocene / lowermost Pleisto-

cene. [Note: This locality was mistakenly referred to as

Morro Abra de los Chaparrinos (the “Huacllaco section”

of DV 1254) in DeVries (2005a, b). |

DV 1252-1 Quebrada de la Vaca, roadeut along Pan-

american Highway, south of south wall, uppermost ter-

race above non-marine de posits, 15°48'56" S$, 74°15'50"

’ (GPS; Chala 1:100,000 quadrangle), uppermost

eae

DV 1254-Bal 6 Section along Panamerican Highway,

ten km southeast of Chala and above Playa Huacllaco. 35

meters above basement rocks in me: | section (see

Figure 3), 15°53'25" S$, 74°09'52" W (GPS: Chala 1:

100,000 quadrangle), upper lower Se me.

DV 1254-Bal 8 Section along Panamerican Highway,

ten km southeast of Chala and above Playa Huacllaco. 42

meters above base see rocks in measured section (see

Figure 3). 15°5 53/25" S, 74°09'52" W (GPS: Chala 1:

100,000 quadrangle). Upper Pliocene.

DV 1254-Bal 10 Section along Panamerican Highway,

ten km southeast of Chala and above Playa Huacllaco.

47.5 meters above basement rocks in measured section

(see Figure 3), 15°53'25" S$, 74°09'52" W (GPS; Chala

1:100,000 quadrangle), upper Pliocene.

DV 1598-1 Roadcut along Panamerican pee de-

scent from north into Yauca. Shell beds. 15°39/49" S,

74°31'50" W (Yauca 1:100,000 nace . lower

Pliocene.

DV 1629-1 Section above Playa Huacllaco, ten km

southeast of Chala, uppermost ben es aie (Unit IV;

see Figure 3), 15°52'47" S, 74°10'13" W (GPS; Chala

1:100,000 quadrangle). Uppermost eae nee

Pleistocene.

DV 1635-1 Yauca Depression, west of Panamerican

Highway. 15°39'33" S$, 75°34'54" W (GPS, Yauca 1:

100,000 quadrangle), lower Pliocene.

THE NAUTILUS 120(4):150-155, 2006

Page 150

A new species of Hoplodoris Bergh, 1880 (Gastropoda:

Opisthobranchia: Nudibranchia) from the Atlantic Ocean

Marta Dominguez

Area de Biologia Animal

Facultad de Ciencias del Mar

Universidad de Vigo

Lagoas-Marcosende, 36310 Vigo

SPAIN

[email protected]

SPAIN

[email protected]

Francisco J. Garcia

Departamento de Fisiologia y Zoologia

Facultad Biologia; Unive said de Sevilla

Apartado 1095, 41080 Sevilla

Jesus S. Troncoso

Area de Biologia Animal

Facultad de Ciencias del Mar

Universidad de Vigo

Lagoas-Marcosende, 36310 Vigo

SPAIN

[email protected]

ABSTRACT

Discodoridid nudibranchs belonging to the genus Hoplodoris

Bergh, 1880 have not been recorde a previously from Atlantic

waters. In the present paper a new species found in Iha de

Cabo Frio (¢

Rio de Janeiro), is described and illustrated. The new species is

seas ed with other species of Hoplodoris. The new species

can be distinguished from other species by its distinctive body

color pattern and by characters of the reproductive system and

radular morphology.

Additional Keywords: Brazil, southwestern Atlantic, sea slug,

Discodorididae, Hoplodoris hansrosaorum.

INTRODUCTION

Up to now, seven species of Hoplodoris Bergh, 1880 are

known (Fahey and Gosliner, 2003). Some species had

been see assigned to the genus Carminodoris

Bergh, 1889. The original descriptions of the genera

Hoplodoris and Carinonons were done by Be roh ( (in

1880 and 1889, respectively). However, the incomple te

de scriptions of type species and the lack of additional

specimens since the original descriptions are some of the

reasons for the difficulties in undertaking taxonomic re-

search in this group ( (Fahey and Cocince 2003). Since

the 1880s, different authors have compiled a list of the

distinguishing characters of Hoplodoris (Thompson,

1975: Miller, 1991: Valdés, 2002). Fahey and Gosliner

(2003) examined discodorid specimens from the type lo-

calities and concluded that the genus Carminodoris is a

junior synonym of Hoplodoris; however, later, Dayrat

ancl Gosline ry (2005) retained both genera,

The genus Hoplodoris has thus far only been found in

the Indian and Pacific Oceans. The present study is the

first record of Ho} jlodoris from the Atlantic Ocean.

An Mbrevetion used in the text is MZUSP, for Museu

de Zoologia da Universidade de Sao Paulo (Brazil).

Cabo Frio Island), Brazil (Arraial do Cabo, State of

SYSTEMATICS

Family Discodorididae Bergh, 1891

Genus Hoplodoris Bergh, 1880

Hoplodoris hansrosaorum new species

(Figures 1-14)

Description: EXTERNAL ANATOMY (FIGURES |, 2): Liv-

ing animal measuring 27 mm length. Body oval, mantle

covered with rounder d tubercles. “Colomicon on central

dorsum of living animal orange-brown, becoming whit-

ish-orange toward mantle edge. Rounded orange-brown

spots around median dorsum, among rounded tubercles.

Opaque white ring present at base of tubercles, tubercle

tip pale-orange. Rhinophore s orange, terminally with

sae cream tip, rhinophores with approximately 15

lamellae. Branchial leaves six, tripinnate. Two anterior

leaves yellowish-cream, two posterior leaves orange. Me-

dian leaves with orange base, upper half vellowien:

cream. Foot narrow and elongated, with notch on ante-

rior side. Oral tentacles two, short. Color of mantle un-

derside whitish, with several brown orange spots on each

side of foot.

BuccAL ARMATURE (FicuRES 3-13): Radular formula

24%31.0.31. Lateral teeth a with denticles

along outer margin of cusp (Figure 3, 9). Middle radular

teeth with a more elongate cusp ce eae ately 14

denticles. Inner side of cusp devoid of denticles (Figure

S). Approximately eight outermost lateral teeth usually

flat, shaped as e sJongate plates; plates closely packed to-

gether (Figure 10, 11). Each plate surrounded by very

fine denticles, plates becoming smaller toward external

side of radula (Figures 4, 12). Labial cuticle with elon-

gate jaw rodlets, some rodlets with rounded tips and

other irregularly tipped (Figures 5, 13)

REPRODUCTIVE SYSTEM (FIGURES 6, a 14): Bulbous

ampulla folded into itself. Prostate rounded, lying next to

ampulli i, narrowing into deferent duct. De fe rent duct

widening into penial bulb. Bursa copulatrix large, ovate.

M. Dominguez et al., 2006

Figure 1-2. Hoplodoris hansrosaorum. Living holotype, 27 mm length. 1. Dorsal view. 2. Ventral view.

Ss } g g

R |

ae |

A ye /\ ,'

¢ a

Figures 3-7. Hoplodoris hansrosaorum. 3. Middle lateral tooth showing the denticles. 4. The three outer lateral teeth. 5. Jaw

rodlets. 6. Reproductive system (ag: accessory gland; amp: arnpulla; be: bursa copulatrix; dd: deferent duct; fg: female gland; ga:

genital atrium: hd: hermaphrodite duct; pb: penial bulb; pr: prostate; rs: receptaculum seminis; st: stylet; vd: vaginal duct). 7. Detail

ot the accessory gland duct with the stylet. Scale lines (3-5) = 10 pm; (6) = 0.5 mm

Page 152

THE NAUTILUS, Vol. 120, No. 4

Figures 8-14. Hoplodoris hansrosaorum. 8. Inner lateral teeth, showing the inner margin of the cusp. 9. Middle lateral teeth.

10-12. Outer lateral teeth, 13. Jaw rodlets. 14. Stylet with broken tip. Scale lines (8, 9, 11, 13) = 20 pm; (10, 14) = 50 pm;

12 10 yum

3ursa about three times as large as receptaculum semi-

nis. Vaginal duct emerging from base of bursa, thick and

looped (Dissection of the only available specimen did

not reveal the presence of genital armature.) Large, elon-

gate accessory sland present, opening into genital

atrium. Gland has narrow duct armed with long, tapering

stylet (Figures 7, 14).

S. Troncoso and Francisco J. Garcia, 25 July 1999

Type Locality: | Collected in the intertidal zone at Hha

de Cabo Frio (Cabo Frio Island), Arraial do Cabo, State

of Rio de Janeiro, Brazil.

Etymology: — The species is named after Hans Bertsch,

ereat friend and malacologist, and his wife Rosa.

DISCUSSION

Several characters allocate the species into the genus

Hoplodoris: species in this genus are characterized by

M. Domimeguez et al.,

2006

Page 153

Table 1. Diagnostic characters in species of Hoplodoris.

H. hansrosaorum

H. armata

H. bifurcata

H. bramale

References

Distribution

Ground color of

dorsum

Tubercle color

Tubercles

morphology

Rhinophore color

Branchial leaf

color

Radula

Jaw rodlets

Accessory gland

Receptaculum

seminis

Ampulla

Dominguez et al.

(present study)

Brazil

Brown orange central

dorsum, and whitish

orange towards the

mantle edge

Brown orange, with a

white ring at the base

Rounded

Orange with cream tip

Anterior leaves, yellow

and middle and

posterior leaves orange

Formula: 24 x 31.0.31.

Middle lateral teeth

with about 14

denticles. The eight

outermost lateral teeth

are elongated plates

with denticles. No

hamate.

With rounded and

irregular tip

One long large gland

Much smaller than the

bursa copulatrix

Bulbous, doubled on

itself

Baba, 1993

Japan

Yellowish gray with some

brown flecks

Brown with a white ring

basally. Some

specimens promine nity

capped with chocolate

brown

Rounded of different

sizes

Dark yellowish gray

Darker yellowish gray

leaves

Formula; 45 x

50—60.0.50—60.

Smooth teeth

Fibrous

Two glands

Small and elongate

Elongated

Fahey and Gosliner,

2003

Hawaii, Japan,

Philippines and

Okinawa

Gray with mottled

shades of darker grays

and reddish-

browns. Central

red-brown band

With a white ring at the

base, followed by a

reddish-brown or tan

ring, and a white tip

Large, round and

tapered

Tan color with white tip

Tan with whitish flecks

the anterior leaves.

Red-brown the

posterior

Formula: 25 « 31.0.31,

Innermost lateral tooth

bifid. Middle radular

teeth with 5-8

denticles. Two or

three outermost teeth

shorter and denticlate

Irregularly tipped

One large and bulbous

gland

Much smaller than the

bursa copulatrix

Long, thin and tubular

Fahey and Gosliner,

2003

Pacific Coast of Costa

Rica

Brown or tan

Brown with a white ring

at the base

Large and rounded

Brown with a white tip

Light tan

Formula: 28 x 38.0.38.

Innermost and middle

teeth smooth. Two

outermost teeth

smaller and

denticulate

Irregularly tipped and

with irregular edges

Two bulbous glands

About the same size as

the bursa copulatrix

Long, thin and tubular

presence of simple, rounded tubercles covering the dor-

sum, head with two conical oral tentacles, anterior bor-

der of the foot grooved and notched, labial armature

armed with jaw elements, radula composed of denticu-

late hamate teeth, and reproductive system with a large

and eng accessory gland ar med with copulatory

spine (Valdés, 2002).

Hoplodoris hansrosaorum new species is externally

similar to H. bramale Fahey and Gosliner, 2003, and H.

grandiflora (Pease, 1860) from Indo-Pacific. The

mantles of both H. bramale and H. grandiflora are brown

and have tubercles surrounded by a white ring, but H.

hansrosaorum has a brown-orange central domuni and is

white-orange toward the mantle edge. Hoplodoris bra-

male and H. grandiflora have tan or brown rhinophores

and gill leaves, whereas the rhinophores of H. hansro-

saorum are orange and the gill leaves are cream and

Orange.

Internally, H. hansrosaorum differs from H. bramale,

particularly in relation to their radular teeth. Hoplodoris

hansrosaorum has teeth with denticles on the outer mar-

gin, and jaw elements with rounded tip, whereas H. bra-

male has teeth devoid of denticles, except for the two

outermost ones; the jaw rodlets are irregularly tipped and

tem also distinguish the two species. Hoplodoris bi -amale

has long and tabular ampulla, two accessory glands, and

the receptaculum seminis is about the same size as the

bursa copulatrix. However, H. hansrosaorum has a thick

ampulla, one accessory gland, and the receptaculum

seminis is about a third the size of the bursa copulatrix.

The radular teeth of H. hansrosaorum and H. grandi-

flora are similar. Both species have lateral teeth with up

to 14 denticles on the outer edge of the cusp, and the

outer lateral teeth have small denticles. Hoplodoris hans-

rosaorum has approximately eight outermost lateral teeth

as plate s, which are very eee to each other, and each

plate is surrounded by very fine denticles. However, H

Page 154

THE NAUTILUS, Vol. 120, No. 4

Table 1. Continued.

H. estrelyado H. flammea

H. grandiflora H. nodulosa

Gosliner and Behrens, 1998;

Fahey and Gosliner, 2003

Philippine Islands, Western

Australia, Vietnam,

Indonesia, Marshall

Islands, Solomon Islands,

Coral Sea

Tan with brown and white

specks

Fahey and Gosliner, 2003

Indonesia

Reddish-mottled brown or

tan. Bright red central

dorsum, with white areas

around them

Bright red in the center of

the dorsum, with white

ring at the base. Reddish-

brown tubercles along the

mantle edge, some of

them with white tip.

Large and rounded

White patch with yellow

center, surrounds a

medial group of brown

tubercles. Some white,

and tan tubercles.

Large tapering tubercles

Fahey and Gosliner, 2003 Fahey and Gosliner, 2003

Eastern Australia, Western

Australia and

New Zealand

Hawaii, Philippines,

Tanzania, Palau,

Mauritius, Kerama Island

and Madagascar

Gray to yellows and

reddish-browns, with

mottled shades of darker

hues. Color more distinct

on the central dorsum

On the central dorsum the

tubercles have the same

coloration of mantle.

Tubercles with white ring

at base towards the

mantle edge

Large, round and tapered

Brown or tan. Some

specimens with dark spots

near the mantle edge.

Others with a dark ring of

tubercles on dorsum

Some specimens with areas

of white tubercles as

perpendicular rays along

the mantle edge

Large and rounded

Reddish brown with white

specks

Tan with brown and white

specks

Formula: 26 x 65.0.65,

Innermost lateral teeth

with two denticles on

outer edge. Middle teeth

with up to 11 denticles.

Outermost teeth with

6-11 denticles

[rregularly tipped

One very large irregularly-

shape od sland

Smaller than the bursa

copulatrix

Thick and tubular

Tan and brown with white

tip

Light gray with tan tips

Formula: 31 x 50.0.50.

Innermost lateral tooth

with a secondary cusp.

Middle lateral teeth with

about 8 denticles. Five

outermost teeth small and

denticulate

Irregularly tipped and with

knurls along the length

One bulbous ‘gland

About half the size of the

bursa copulatrix

Long, thin and tubular

Light tan with white tip

Light tan with frosted tips

Formula: 26-40 x

40-116.0.116—-40.

Innermost lateral teeth

with or without denticles.

Middle lateral teeth with

up 14 denticles. Two or

three outermost teeth

hamate with denticles.

Irregularly tipped

One long and tubular gland

Much smaller than the

bursa copulatrix

Thick and tubular

Tan with a white tip

Tan anterior leaves with

whitish flecks. The

posterior leaves can be a

darker color

Formula: 25 x 53.0.53,

Smooth innermost and

middle teeth. Four

outermost lateral teeth

short and denticulate

Irregularly tipped

Two long and bulbous

elands

Much smaller than the

bursa copulatrix

Long and tubular

grandiflora has outer lateral teeth with small denticles,

and the two or three penultimate lateral teeth are hamate

and much shorter than the rest. The reproductive system

of the two species are similar, but the ampulla and the

accessory gland of H. grandiflora are long and tubular

(bulbous in H. hansrosaorum).

There are differences between Hoplodoris hansro-

saorum and other Hoplodoris species. For example, the

coloration of Hoplodoris bifurcata (Baba, 1993) is com-

Fahey and Gosliner, 2003). The

bac koround color in this latter specie Ss is gray with

plex and variegated (

mottled shades of darker grays and reddish- browns scat-

tered over the mantle, whereas H. hansrosaorum has

whitish-orange background color. Hoplodoris bifurcata

has a red-brown band of color band on the central dor-

sum, which is lacking in H. hansrosaorum. Both species

have a white ring at the base of the dorsal tubercles, but

H. bifurcata has also a red-brown or tan ring on the

tubercles and a white tip. Hoplodoris bifurcata has tan

and white rhinophores, the anterior gill leaves are tan

with whitish flecks of color, and the posterior leaves are

red-brown. This coloration differs from that of H. hans-

rosaorum, because the rhinophores are orange with a

pale cream tip, and the gill leaves are yellow-cream and

tee Further, the “aduler morphology differs between

these species. T hei imermost lateral tooth of H. bifurcata

is bifid, the middle teeth have a cusp with 5-8 denticles,

and the 2—3 outermost lateral teeth are denticulate

plates. Hoplodoris hansrosaorum has an innermost lat-

eral tooth with one cusp, the middle teeth have a cusp

with about 14 denticles, and the eight outermost lateral

teeth are denticulate plates.

M. Dominguez et al., 2006

The background color of Hoplodoris nodulosa (Angas,

1864) ranges from gray to yellow and reddish- es a

white ring may be present at the base on the tubercles.

That species differs from H. hansrosaorum by its tan-

colored rhinophores and tan gill leaves with whitish

flecks. Furthermore, the four outermost radular teeth of

H. nodulosa are denticulate (the rest are smooth), and

there are two accessory glands in the reproductive sys-

tem.

Main internal and external morphological characters

that can be used to distinguish among the species of

Hoplodoris are compiled in Table 1

In relation to geographic distribution, the species -

grandiflora, H. bifurcata, H. nodulosa, H. bramale,

estrelyado Gosliner and Behrens, 1998, H. flammea ee

hey and Gosliner, 2003, and H. armata (Baba, 1993), are

known only from the Pacific and ee Ocean. There-

fore, H. hansrosaorum is the first record of Hoplodoris

from the Atlantic Ocean.

ACKNOWLEDGMENTS

The authors are very grateful to Dr. Paulo Marcio Costa

and Dr. Renata Gomes for assistance in collecting nudi-

branchs; the Rear-Admiral

the Instituto de Estudos do Mar of the Brazilian Navy,

for facilitating the transport by boat in the zone of Arraial

do Cabo, and Jestis Méndez for the scanning electron

micrographs at CACTI of Vigo University. This paper

Napoleaio Gomes, director of

has been partially supported by the projects PHB2002-

0045-PC and CGL2004-20366-E/BOS, of the Ministerio

de Educacion y Ciencia (MEC, Spain).

LITERATURE CITED

Bergh, R. 1880. Malacologische Untersuchungen. Theil 4,

Suppl. 2. In: C. Semper (ed.) Reisen im Archipel der Phil-

ippinen. Kreidel, Wiesbaden, pp. 51-56.

Bergh, R. 1889. Malacologische Untersuchungen. Theil 3, Heft

16. In: C. Semper (ed.), Reisen im Archipel der Philip-

pinen. Kreidel, Wiesbaden, pp. $15-872, pls. 82-84

Dayrat, B. and T. M. Gosliner. 2005. Species names and meta-

phyly: a case study in Discodorididae (Mollusca, Gas-

tropoda, E uthyneura, Nudibranchia, Doridina). Zoological

Scripta 34; 199-224.

Fahey, S. J. and Gosliner, T. M. 2003. Mistaken identities: On

the Discodorididae ae Hoplodoris Bergh, 1880 and

Carminodoris Bergh, 1889 (Opisthobranchia, Nudibran-

chia). Proceedings of the California Academy of Sciences

54(10): 169-208.

Miller, M. C. 1991. On the identity of the dorid nudibranch

Homoiodoris novaezelandiae Bergh, 1904 (Gastropoda:

Opisthobranchia). Journal of Natural History 25; 293-304.

Thompson, T. E. 1975. Dorid nudibranchs from eastern Aus-

tralia ( (Gastropoda, Opisthobranchia). Journal of Zoology

176: 477-517.

Valdés, A. 2002. A phylogenetic analysis and systematic revision

of the cryptobranch dorids ( Mollusca, Nudibranchia, An-

thobranchia). Zoological Journal of the Linnean Society

136: 535-636.

THE NAUTILUS 120(4):156-161, 2006

Page 156

First report of the invasive freshwater snail Tarebia granifera

(Lamarck, 1816) (Gastropoda: Thiaridae) from Israel

Frida Ben-Ami’

Department of Evolution, Systematics and

Ecology

The Hebrew Unive rsity

Jerusalem 91904

ISRAEL

Frida. [email protected]

ABSTRACT

An invasive pda gastropod, the freshwater snail Tarebia

granifera, is reported for the first time from Israel. In a survey

of 52 freshwater sites populations of T. granifera were discov-

ered at four sites, in abundance at two of them, one of which

included individuals infected with the trematode Philophthal-

mus sp. In both sites T. granifera outnumbered its close native

relative and potential competitor, the thiarid Melanoides tuber-

culata, in abundance and mean embryo counts, and in one of

the habitats almost exchided it. This invasion may have impor-

tant public health implications, due to T. granifera’s role as an

intermediate host of the human eye flukes. Furthermore, evi-

dence of long-distance dispersal or multiple invasion sites may

facilitate the host’s and parasite’s spread to other water bodies

used for agriculture and fish ponds.

Additional Keywords: Melanoides tuberculata, Philophthalmus,

biological invasion.

INTRODUCTION

Biological invasions threaten terrestrial, marine and

fre shivate ry biodiversity, as increased replacement of na-

tive s by highly competitive generalist invaders

may lead to biotic homeee nization (Prenter et al., 2004).

In particul: uw, agricultur al trade has broken down many

natural disperse ul barriers (Kolar and Lodge, 2001) and

facilitated the invasions of many exotic species, with dra-

matic effects on ecosystem functioning (¢ Carlsson, Bron-

mark, and Hansson, 2004). Among the most notorious

invaders are mollusks, which cause extensive economic

loss and disturbance because of their detrimental impact

on indigenous fauna and their intermediary role in trans-

mitting parasitic diseases to humans (Pointier, 2001).

\mong gastropods, the freshwater snail Melanoides tu-

herculata (Miiller, 1774), which is native to the Middle

1 , +

Present address: Zoologisches Institut Evolutionsbiologie

Universitit Basel, Vesalgasse 1, 4051 Basel, Switzerland

East (Pointier, 1999), has invaded North America (Mur-

ray, 1964; Murray, 1971), South America (Pointier et al.,

1994; De Marco, 1999; Giovanelli, Vieira, and da Silva,

2005), East Africa (Genner et al., 2004), the French

Polynesia (Pointier and Marquet, 1990) and the islands

of the Pacific (Cowie, 2001b).

Another invasive freshwater snail is Tarebia granifera

(Lamarck, 1816) (Thiaridae). It is native to Southeast

Asia, from India and Ceylon eastward to the Philippines

and Hawaii, northward to southern Japan, and southward

to the Society Islands (Abbott, 1952), where it is found in

a wide variety of habitats including rivers, streams, lakes,

drainage ditches, irrigation canals, cement ponds and

swamps (Chaniotis et al., 19SOb), though shallow, fast-

flowing freshwater streams seem to be its pre ferred habi-

tat (Lachner, Robins, and Courtenay, L970). Tarebia

granifera can survive in a wide range of temperatures

(740°C), can be kept out of water for several days under

relative humidity of 80-100% (Chaniotis et al., 1980c)

and is even capable of surviving in polluted habitats

(Chaniotis et al., 1980b). It feeds on algae, associated

microorganisms and small particles of organic matter

(Ogle soy, 1977). Tarebia granifera is ovoviviparous and

reproduction is mainly by parthenoge nesis (Morrison,

1954), although male presence varies from total absence

in populations from Florida and Guam (Abbott, 1952) to

frequencies of 22.7% in Puerto Rico (Chaniotis et al.,

19S0a).

Tarebia granifera serves as an intermediate host of

numerous trematodes that cause parasitic diseases in hu-

mans. Among these are representatives of the superfam-

ily Hete rophyid: ve, Which are vectors of intestinal flukes

of animals and humans, e.g., Metagonimus yokogawai

(Murray and Stewart, 1968). Other trematodes include

Philophthalmus megalurus (family Philophthalmidae )

which affects the nictating membrane of aqui atic birds

(Jacobson, 1975). As part of the life cycle of the trema-

todes, a se onda intermediate host, such as a freshwater

crayfish or crab, is necessary and for human infection to

be successful, the infected crustacean must be eaten raw

(Lachner et al., 1970).

F. Ben-Ami, 2006

This study documents the first record of T. granifera

from Israel and discusses the possible implications of its

invasion on public health and on the native freshwater

molluscan fauna.

MATERIALS AND METHODS

A survey of 52 freshwater sites was conducted in Israel

during July-August 2004 (Figure 1). At each site five

random samples were taken by dredging with a square-

framed net (50x50 cm, 1 mm mesh) and each sz unpled

scanned for the snails retrieved. Additionally, at two sites

that included T. granifera, all snail species found within

twenty random squares (20x20 em) were collected and

shell height was measured to an accuracy of 0.01 mim.

Most snails were returned to the site. However, to quan-

tity embrvo numbers and infection prevalenc e, LOO ran-

domly chosen individuals of T. granifera and M. tuber-

culata were tr: insported alive to the laboratory. The num-

ber of embryos in each brood pouch was counted and

trematode infection determined by examination of the

gonad and digestive gland under a light microscope. In-

dividual snails were defined as adults if they measured

more than 7 mm in shell height and treated as juveniles

if their height did not exceed 7 mm. This definition is

based upon the dissection of 266 snails in which embryo

presence was scored against shell height.

For statistical analysis, SPSS for Windows version

12.0.2 (SPSS Inc., 2004) was used. Means +SD were

determined and all probabilities discussed are two- tailed.

Frequency data were analyzed either with the x~ test or

Fisher's exact test, when possible. An analysis of covari-

ance (ANCOVA) was used to compare mean embryo

counts among habitats and species (see Table 1 for de-

tails).

RESULTS

Tarebia granifera was only found at four of the 52 sites,

all in the Jordan Valley and within a radius of 1.5 km: En

Tayuon, En um Sidra, En Saharon and Avoka ( Figure 1).

Since only three T. granifera individuals were found in

En Tayuon and two individuals in En um Sidra, no fur-

ther studies were undertaken beyond recording these

observations. In En Saharon (n=1386) and Avoka

(n=1740) four relatively abundant snail species were

found: T. granifera, M. tuberculata, Melanopsis saulcyi

(Bourguignat, aoe ) and Theodoxus jordani ( (Sowerby,

1836) (Figure 2

which only a fow individuals were found, Heleobaia con-

tempta (Dautzenberg, 1894) and Melanopsis buccinoidea

Olivier. 1901).

Species abundances diff a significantly between En

Saharon and Avoka (~=389.3 df=5; P<0.001). Specifi-

cally, T. granifera (Fisher's exact test; P<0.001), M. buc-

cinoidea (Fisher's exact test; P=0.004), T. jordani (Fish-

. Additionally, there were two species of

Mediterranean Sea

ne}

Figure 1 ee of sampling sites in Israel: (1) En

Tayuon, (2) Avoka, (3) En um Sidra and (4) En Saharon. Due to

the een of ie map, some data points represent more

than one sampling site.

er’s exact test; P<0.001) and H. contempta ( Fisher's exact

test; P=0.006) were significantly more abundant in

Avoka, whereas M. nibereuaia (Fisher’s exact test:

P<0.001) and M. saulcyi (Fisher's exact test: P<0.001)

were more common in En Saharon.

Page 158

THE NAUTILUS, Vol. 120, No. 4

50 TDEn Saharon

BAvoka

te}

o4 ee ee Ep ms _ _—__

Figure 2. Relative species abundances in En Saharon and

Avoka.

The size distribution of T. granifera in En Saharon was

highly bimodal, with the majority of individuals in the

sample being young (56.5%) and with an average shell

height of 2 8414 min, compared to 43.5% of adults with

mean shell height of 18.2+3.4 mm (Figure 3A). In Avoka,

however, size distribution was unimodal and skewed to-

wards adults (78.8%; average height of 10.6+2.5 mm), in

contrast to 21.2% of young wath mean shell height of

5.0+1.5 mm (Figure 3B).

The size distribution of M. tuberculata in both sites

was unimodal and skewed towards adults. The En Sa-

haron sample consisted of 74.1% of adults with an aver-

120

Frequency

co 8 6&6 8 8 8

Frequency

b for) for) S DS

o lU8 o.6hlUS 8S Ss 8

15 25 30

Height (mm)

Figure 3. Size distribution (mm) of Tarebia granifera in (A)

En Saharon and (B) Avoka

age shell height of 12.6+3.4 mm and 25.9% of young with

mean shell height of 4.2+1.8 mm (Figure 4A). Similarly,

in Avoka there were 67.9% of adults with an average shell

height of $.9+1.7 mm and 32.1% of young with mean

shell height of 5.8+0.8 mm (Figure 4B).

The juvenile-adult ratio differed significantly bets tween

T. granifera and M. tuberculata in En Saharon (y7=56.0:

df=1; P<0.001) but not in Avoka (x7=1.9; df=1; P=0.165).

Mean embryo counts of T. tre were 28.1+21.5

(n=100) in En Saharon and 9.6+9.3 (n=100) in Avoka.

The mean number of embryos per M. tuberculata female

was 20.5+17.6 (n=101) in En Saharon and 1.1+1.7 (n=7)

in Avoka. Mean embryo counts were significantly higher

in En Saharon than in Avoka for both tae, but did not

differ significantly between species (Table 1

Inf fected snails were found only in En ake The

brood pouch of six T. granifera individuals (6%) was

infected by both miracidia and cercariae stages of

Philophthalmus sp. One M. tuberculata individual (1%)

was infected by Centrocestus sp. In both species infected

snails were not carrying embryos.

The National Mollusc collection of Israel has no

records of T. granifera trom Israel. It is also absent from

the rich fossil ane of the Jordan Valley, currently

under investigation (J. Heller, personal communication).

DISCUSSION

This is the first ig aa of T. granifera from Israel, where

it was found in four freshwater springs in the Jordan

Frequency

roy

3 15

10 4

B 0 5 10 15 20 25 30

Height (mm)

Figure 4. Size distribution (mm) of Melanoides tuberculata

in (A) En Saharon and (B) Avoka.

F. Ben-Ami, 2006

Table 1. Analysis of covariance testing the effects on female

fecundity (number of embryos), with habitat and species as

fixed factors and shell he ight as covariate (R° = 0.463). Embryo

counts were Box-Cox transformed prior to analysis to meet the

normality assumption, The homogeneity of variances criteria

was also met using Levene’s test of equality of error variances

F = 2.6; df, . = 3262; P = 0.054.

Sum of

Source dt squares F-ratio P

Whole model 4 23585.5 56.3 <0.001

Shell height ] 9142.4 87.3 <0.001

Habitat | 1587.6 15.2 <0.001

Species 1 34.5 0.3 ns

Habitat x Species | 213.6 2.0 ns

Error 261 27344.6

Valley. The mode of introduction is unknown but it is

likely that, as in Martinique (Pointier et al., 1998), T.

granifera arrived via the trade in aquatic plants and pet

fish. The invasion of T. granifera into Israel poses an

ecological threat to native freshwater mollusks. In par-

ticular, granifera is likely to compete with other

thiarids, notably M. tuberculata (Pointier et al., 1998), as

these two species are similar in form and occupy a closely

similar microhabitat of soft mud substratum.

The density of T. granifera in Avoka (1139 snails/m?;

52.4%) was more than twice that in En Saharon (511

snails/m7: 29.5%), whereas the density of M. tuberculata

in Avoka (35 snails/m*: 1.6%) was almost tenfold less

than in En Saharon (290 snails/m?: 16.7%). This may

suggest that in sites where T. granifera reaches high den-

sities, it competitively excludes M. tuberculata. The bi-

modal size distribution of the T. granifera population in

En Saharon, consisting of adults and very young indi-

viduals, may suggest that in this site T. granife ra is “still in

the midst of the establishment process. It could perhaps

indicate that the life-history of T. granifera is different,

ie. adults breed only at a certain time of year and the

small size class represents a new cohort of young snails.

If that is the case, since M. tuberculata reproduces

throughout the year (Ben-Ami and Heller, in press), the

T. granifera population must compensate for shorter

breeding periods via increased reproduction or reduced

juv enile mortality. This may in turn explain the relative

success of the M. tuberculata population in En Saharon,

whereby the population structure represents the stage in

which both species can share the habitat, without com-

petitively excluding each other. Although only a long-

term survey of multiple invasion sites in Israel can un-

ravel the exact nature of the competition between the

two species, a similar situation was observed in Martin-

ique, where T. granifera quickly outnumbered M. tuber-

culata, though it did not exclude it entirely (Pointier et

al., 1998S).

The mean number of embryos was larger (though not

significantly) in T. granifera than M. tuberculata females

at both sites, which in part may explain the competitive

advantage of T. granifera over M. tuberculata. Alterna-

tive nonexclusive factors, which may explain the com-

petitive advantage of T. granifera, inc ‘ude higher survival

rates of juveniles or lower susceptibility to predators,

although the latter is less likely because there are no

spe cific molluscivores in either site. Additionally, my

field observations reveal that M. tuberculata occurs only

along the banks of rapidly flowing rivers, whereas T.

granifera is distributed over the entire river bed, sug-

gesting that the latter specie s can withstand faster cur-

rents and resist dislodgement.

The invasion of T. granife ra into Israel may also influ-

ence public health, as it was found to be infected by

philophthalmid eye flukes. These parasites are known in

Israel from both M. tuberculata, which hosts Philoph-

thalmus distomatosa (Radev, Kanev, and Gold, 2000)

and M. buccinoidea, which hosts P. palpebrarum (Gold,

Lang, and Lengy, 1993). Although a case of human in-

fection by P. palpe -brarum in isvaal was documented al-

most thirteen years ago (Lang et t al., 1993), the genus

Philophthalmus appears to be very rare and has not been

found in Israel since then (D. Gold, personal communi-

cation). Given that six T. granifera individuals (6%) of

our sample were found to be infected, and that the para-

site's lifecycle requires only two hosts (a snail, and a

bird or mammal), this may suggest that Philophthalmus

could spread relatively easily. However, in this study M.

tuberculata and Me lanopsis ‘buccinoidea were not found

to be infected with Philophthalmus. Since T. granifera

has been found to host Philophthalmus megalurus

in Cuba (Jacobson, 1975), it may very well be that the

T. granifera population found in Israel is infected

with a new or another Philophthalmus species, the

spread of which depends solely on the invasion success of

T. granifera. This may in turn also affect the parasite

fauna in the Jordan Valley (Torchin, Byers, and Huspeni,

2005).

Tarebia granifera has a long history of invasion. It was

introduced to Florida and Texas in the 1940s via aquatic

plants (Abbott, 1952; Murray, 1964; Murray and Wop-

hall 1965). It was recorded from Puerto Rico by Harry

and Aldrich (1958) and then spread quickly throughout

the Caribbean, reaching the Dominican Republic by

1967 and Cuba, Grenada and Venezuela in 1970 (Jaume,

1972; Chrosciechowski, 1973; Jacobson, 1975; Ferguson,

1977; Pointier et al., 1994). In 1991 T. granifera was

discovered in the Charpentier River, Martinique, where

it rapidly spread from its introduction site and had al-

ready colonized thirteen river systems by 1997 (Pointier

et al., 1998). In places where it invaded it quickly became

dominant in numbers and competed with native Mol-

lusks (Murray, 1971; Jacobson, 1975; Oglesby, 1977;

Chaniotis et al., 1980b). It has also been suggested that T.

granifera is responsible for the disappearance of the en-

demic freshwater snail Hemisinus cubanianus

(d’Orbigny, 1841) in Cuba (Pointier, 1999). A recent re-

port from South Africa suggests that T. granifera has

been present in a reservoir since at least 1996 ( Appleton

and Nadasan, 2002).

Page 160

THE NAUTILUS, Vol. 120, No. 4

Despite its invasiveness or perhaps because of the

competitive capabilities of T. granifera, it has been used

as a biological control agent against the snail hosts of

suhisiosommacix basin Aabrata (Say, 1S1S) and

B. straminea (Dunker, 1848) (Butler et al., 1980; Mich-

elson, 1992; Pointier, 2001). Gomez et al. (1990) sug-

gested that T. granifera probably produces chemical fac-

tors that reduce the fecundity of B. glabrata. Perez et al.

(1991) found that T. granifera competed with B. glabrata

not for food or vital space but for chemical siibisianiees or

by physical contact with a large number of individuals.

Canete et al. (2004) found a negative association between

T. granifera ce Fossaria cubensis (Pfeiffer, 1839) abun-

dances, but a positive correlation with Pseudosuccinea

columella (Say, 1817) densities, both of which are inter-

mediate ee of the liver fluke Fasciola hepatica. This

only emphasizes that the safety and effectiveness of snails

as biocontrol agents remains questionable (Civeyrel and

Simberloff, 1996: Cowie, 2001a).

The dispersal of an alien freshwater snail species fol-

lowing invasion could occur through two non-exclusive

processes. The first is invasion to a single site, followed

by diffusion into neighboring sites at a rate depending on

the local intrinsic rate of population increase and the

diffusion coefficient (Pointier et al., 1998). The second is

long-distance dispersal by means of an agent of dispersal,

such as birds ( (Boag, 1986). miamicials (Madsen and

Frandsen, 1989) or human activities (Pointier et al.,

1998).

The four sites at which T. granifera was found are

isolated from each other. This suggests the occurrence of

either long-distance dispersal or multiple (separate) in-

vasions, though only genetic studies will be able to dis-

tinguish these. Both sites flow naturally into the Jordan

River, which drains many other water bodies, including

water used for agriculture and fish ponds, and thus could

further spread T. granifera. Hence, the dispersal pat-

terns of T. gi canifera throughout the Jordan Valley are

difficult to predict and only a long-term survey can esti-

mate the impact of the invasion.

ACKNOWLEDGMENTS

J. Heller, J-P. Pointier, N. Sivan and an anonymous re-

viewer provided valuable comments that improved the

manuscript. Tam grateful to D. Gold for assistance dur-

ing the identification of the parasites, to N. Sivan for

preparing the map, and to R. Ben-Yossef and Y. Sever for

helping with the sampling. Specimens for this study were

collected under permit 6094 from the Israeli Authority

for Nature Reserves and National Parks. This study was

supported by a Horwitz Foundation Fellowship to FBA

and by grant 665/02 of the Israel Science Foundation to

J. Heller.

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California.

THE NAUTILUS 120(4):162, 2006 Page 162

Notice

American AMS 2007—Antwerp, Belgium

Malacological

Society 15-20 July 2007

fs Antwerp, Belgium

‘A 15 - 20 July 2007

The American Malacological Society (AMS) will hold its 73 annual meeting in Antwerp in July 2007 as part of the next World

Congress of Malacology. Sessions of contributed papers and posters are planned as usual, as are six exciting symposia:

® “Molluscan models: advancing our understanding of the eye” organized by Jeanne Serb and Laura Robles [sponsored by AMS]

® “Sexual selection” (Ronald Chase and Joris Koene)

® “Micromolluses” (Daniel Geiger)

@ “Molluscs as models in evolutionary biology” (Matthias Glaubrecht and Thomas von Rintelen)

® “Neogastropod origins and evolution” (Jerry Harasewych)

Our venue is the Groenenborger Campus of University of Antwerp (UA), hosted by Thierry Backeljau, President of Unitas

Malacologica (UM). Campus facilities include most of our meeting rooms and a cafeteria under one roof. The conference begins with

a Sunday afternoon icebreaker. Scientific presentations are in four parallel sessions on Monday, Tuesday, Thursday, and Friday.

Wednesday is open for you to join one of the suggested tours, discover historical Antwerp, or visit a nearby museum collection.

During the Tuesday evening session we will enjoy Belgian food, wine, and beer. A banquet will conclude the Congress on Friday

evening.

On Thursday evening, we will host the traditional AMS Auction of molluscan books and paraphernalia (no specimens) open to all

Congress participants. This annual event raises the necessary funds for our student programs, in addition to providing a fun-filled

evening for all involved (even if you don’t buy anything!). Paul Callomon will return as our auctioneer—this is not to be missed!

Because we are meeting overseas, transporting books and other items for the auction presents us with a challenge. For this reason,

Lam asking each attending member to bring at least one item for the auction in his/her luggage. Please dig deeply and generously—

surely everyone has an unneeded book or something malacologically silly to contribute! Please help AMS present its best face to the

world!

Early registration ends 30 April 2007. Registration for AMS members who are not also UM members is €280 = ~$221 [see your

favorite online currency converter for current rates] (students €160); a discounted price applies to those who are also UM members.

The registration fee includes the abstract volume, event refreshments, and all lunches.

Airfare from the US to Brussels Intemational Airport is relatively affordable. From there, buses and trains to Antwerp are easy and

inexpensive. Dormitory accommodations are offered at UA (200 single and 20 double rooms, each with linens and washbowl but

shared toilets/showers) at €20-27 per person/night, including breakfast. Hotel rooms are available in Antwerp city center, very near

the railway station and bus terminals. Room prices range from €47.5 (singles) to €155 (4 persons) per night, including breakfast. Buses

travel from the city center to campus in about 20 minutes. Alternatively, the organizers recommend a rental bicyele—the vehicle of

choice in Antwerp for young and old alike!

Student travel grants will be offered by both AMS and UM. Watch the AMS (http:/Avww.malacological.org/meetings) and WCM

websites for details and application forms. There will also be student awards at the Congress for best oral and poster presentations.

Full details about the Congress are available at http:/Avww.ucd.ie/zoology/unitas/congress.html#content.

Please join us in Antwerp!

Paula M. Mikkelsen

AMS President

NAUTILUS

Volume 120

2006

AUTHOR INDEX

BEN=AMIL Bi. saisascicedetinctncdaePeieasceveslssewusweia’ snedassoesune 156 COSTROFSKY,, Ml. Tues sic5 sciedelniss oasdeaiae atid eager eghadiaaeendeeesn 106

BOUDREAUS. MILI... x s2net.aauennsnate eaneaeesed wonnes a ssasmemeh ances 34 OSWATIOG Wis Wo vatogolecancrsntsudisoonautameieonsstancusoutanwces 30

BUTKAS; Kio Ji. sia tiniicssanontnia ins cstuisaainecstgana neue vecacanntes 106 PETIT Ris, auch Peasiiusenckaet ctr uaasndescieas ideas uot 79, 112

GOAN Bie Wise daade oattclereaace taeda ammatineaemaiabaialiumyniachacsguse 112 SAWTA Te TAs snuiceAncadunnace Miensdeandcasteds ata ateraeeoatenneieate nee 66

GOES: Bs F..cce2cobtcctncsne. cose nusanss ooesscaiine cua tees 131 DEDBERRY: Gu Re. dacuts caso ccedaen te bee ras eee datenaades ohmewsdavede 39

IDEVRIES “TJs. sestavsdisressaastacaccieestetsanaes aoeines 1O1, 139 DELLANES lo cncastcsassamicewans Rtaue tian Genmad stuns ven visuiugaintant Ws 15

DOMINGUEZ, MM. o.2sci.igcotacsisaveheodsadearadiatiheet ta sacneces 150 SILVA; Gy Ms DA. a 2icgiatacdsexsitanccnbaastatoebenieadarekssiseweeses Sl]

GARGIA, By Je-asiis terenediseesinays coecinwia cote remo teaccseaeenss 150 SLAPCINSKY,. J. seesia hoon dean taaneavaleadestauiacinciiansaerees 119

GEIGER, Ds Tage cde iiss cadacdens sicsnueddasusndeasstesednacaaameaeas’ 45 DOUTRESS, ea: cxacunietescosaennaas a berahaaueneeer satan takeeeds 66

GROWERS: ME. Mis cute Hea ceaeiocsanac tonsca aaa teases ene connects onan 101 DUTRAUSS I. cheneesceonivsecatvob-enssah nundiaadsoonsieedel eons caamentes 94

HARASEWYCH, M. G. oo... cece sseee esses sesetessteeserseees OO, 94 THOMPSON, FG. sisiccssaicntteinis na vicitesecsatensiasciveaes 21, 25

FUBRYZsGCoIM., codscas tances sanaeeinesesoindenevinesaicntnanes esealene ae 52 FRONGOSO, J. S..sscaeentiaasaangscsaad otgnersagansneashaangunertd 150

FLOUART sINe aq tapatoanesol tence se scttaanutnieace con enutietdeetesane (OD WRBINAS IME 520 detetaa tndicsiesa ae nati Poeaiedeihaasandtanavhanedess 101

NGANIDAU:, (Di, (sc siutdaiasson di deaeiasaacteas sae ee inpatients eqns Sl VILVENS, Cy ce ccctes cance ode nines tyenasbeniotsiae oideorndstciadlaawand ene 15

LBAL eis Elan saiaiteecatenag nneeteaseytesianaies gtead ened tise ss 1, 116 WALSH. GRY, crecisaancondssascrencias seomedaentay madeag bene sgaee 131

MALGHUS, IN: dscssacdidssatardsiedat dntaccasdbvomawnoasag eevee S WALTERS, [b.. l sastivcianiacancieansiacmsmie aneniarieesdnvetnwanctis 34

DEBE WO: elves sat 2aied ace tamatn suet A tia tue-cny-fatects saab in nes semana: 25

NEW TAXA PROPOSED IN VOLUME 120 (2006)

GASTROPODA

Hoplodoris hansrosaorum Domingues, Garcia, and Troncoso, 2006, new species (Discodoridae) . 2... 0. ee 150

Humboldtiana corruga Thompson and Mejia, 2006, new species (Humboldtianidae).. 2... 0.0. 25

Humboldtiana iversoni Thompson, 2006, new species (Humboldtianidae) 2... 0. 21

Humboldtiana sylvania Thompson and Mejia, 2006, new species (Humboldtianidae) 2.0. 28

Margarites huloti Vilvens and Sellanes, 2006, new species (Trochidae) 2... 0. ee 19

Muracypraea ormenoi DeVries, Groves, and Urbina, 2006, new species (fossil, Cypraeidae) 2.0... 2 103

Ornopsis dysis Squires and Saul, 2006, new species (fossil, Buccinidae?) 2... 71

Otukaia crustulum Vilvens and Sellanes, 2006, new species (Calliostomatidae) 2... 16

Paryphantopsis misimensis Slapcinsky, 2006, new species (Charopidae) . 2... ee 123

Paryphantopsis vanatinensis Slapcinsky, 2006, new species (Charopidae) pce pen fet a Eocene os aed yo gg ane arenes Give 125

Perrilliata Squires and Saul, 2006, new genus (fossil, Fasciolariidae?) 2... ee 74

Perrilliata califia Squires and Saul, 2006, new species (fossil, Fasciolaridage) 9s aucu 1a ea ine ks oe eee omg ds 74

Prisogaster mcleani DeVries, 2006, new species (fossil, Turbinidae) 2... 145

Prisogaster valenciai DeVries, 2006, new species (fossil, Turbinidae) .. 2. ee ee nee 143

Sasakiconcha Geiger, 2006, new genus (Anatomidae?) 2... 45

Sasakiconcha elegantissima Geiger, 2006, new species (Anatomidae?) 2.2... ee 46

Saxituberosa Squires and Saul, 3006, new genus (fossil, eee uae Barrie alin gE SWI oti ee oo oe a ee 72

Saxituberosa fons Squires and Saul, 2006, new species (fossil, Fasciolariidae?) 2... 0. ee 72

Scaphella carlae Landau and Silva, 2006, new species (fossil, Volutidae) 2... 0. ee 89

Typhisopsis carolskoglundae Houart and Hertz, 2006, new species (Typhinae) 2.2... 0 ee . 5G

a iia ae

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